Tooth impact rate in the song of a shorthorned grasshopper: A parameter carrying specific behavioral information

Summary Receptive females of the gomphocerine grasshopperOmocestus viridulus L. were offered different artificial grasshopper songs in a one- or two-channel test situation. In the computer generated songs the tooth-impact-rate and the amplitude functions of the chirp were varied.The results of the one-channel experiments showed that the grasshoppers preferred signals with normal tooth-impact-rate to signals with half the rate. The amplitude function seems to play a minor role in the recognition process, unless this parameter is changed very radically.The results of the two-channel tests were ambiguous. A third test series was developed to interpret these results. It was found that female grasshoppers once excited by an acceptable signal afterwards responded to former unacceptable signals. Accordingly, if just one of the two signals is able to excite the female, the female may subsequently choose randomly between the signals. If the signals are of different sound levels, the louder is preferred.The shorthorned grasshopper may interpret the tooth-impact-rate as a frequency. It is proposed that the grasshopper may use the four sense cell groups of the ear as a simple sort of a spectral analyzer. These cells are tuned to different frequencies and/or differently tuned.The running Root Mean Square value of a grasshopper signal, its dependence on the applied integration time and its possible relation to perceived energy are reported in an appendix.     

Acoustic pattern recognition in a grasshopper: processing in the time or frequency domain?

Males of the grasshopper Chorthippus biguttulus produce songs which consist of the stereotyped and rhythmic iteration of a sound unit (termed syllable) separated by distinct syllable pauses. Virgin females respond to this signal, and to similar artificial signals, with song phrases of their own. In behavioural experiments the response probability of virgin females can be measured with artificial acoustic stimuli. The stimuli consisted of an amplitude modulated noise the envelope of which was altered. We investigated several hypotheses on the mechanisms of conspecific song recognition with special emphasis on the question whether recognition occurs in the frequency domain or in the time domain. (1) Females of Ch. biguttulus required only the first five Fourier components of the envelope function (corresponding to 50 Hz for a fundamental frequency of 10 Hz) to detect the syllable/pause structure. In addition, they detected small gaps within syllables if the signal contained at least ca. 15 Fourier components (corresponding to a frequency of 150 Hz). Further experiments showed that the correct phase information of the Fourier components is necessary for recognition, indicating that pattern recognition is not achieved merely on the basis of band pass filtering. (2) A cross correlation between the signal and an assumed internal template yields only inconsistent predictions of the response probabilities. (3) The recognizer system probably works in the time domain, possibly by direct comparison of adjacent syllable and pause durations. It is not yet clear whether the duration of a syllable is evaluated with respect to the preceding or succeeding pause. We emphasize that the neural recognizer of the grasshopper does not only examine a signal for its similarity to an internal template, but that it also takes into account features that indicate an incorrect signal. This may be a general feature of neuronal pattern recognition systems which have been shaped by natural selection. Received: 4 October 1997 / Accepted in revised form: 26 August 1998     

A highly sulfated chondroitin sulfate preparation, CS-E, prevents excitatory amino acid-induced neuronal cell death

Chondroitin sulfate (CS) is a major microenvironmental molecule in the CNS, and there have been few reports about its neuroprotective activity. As neuronal cell death by excitotoxicity is a crucial phase in many neuronal diseases, we examined the effect of various CS preparations on neuronal cell death induced by the excitotoxicity of glutamate analogs. CS preparations were added to cultured neurons before and after the administration of glutamate analogs. Then, the extents of both neuronal cell death and survival were estimated. Pre-administration of a highly sulfated CS preparation, CS-E, significantly reduced neuronal cell death induced by not only NMDA but also ( S )-α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid or kainate. Neither CS preparations other than CS-E nor other highly sulfated polysaccharides such as heparin and dextran sulfate exerted any neuroprotective effects. NMDA-induced current in neurons was not changed by pre-administration of CS-E, but the pattern of protein-tyrosine phosphorylation was changed. In addition, the elevation of caspase 3 activity was significantly suppressed in CS-E-treated neurons. These results indicate that CS-E prevents neuronal cell death mediated by various glutamate receptors, and suggest that phosphorylation-related intracellular signals and the suppression of caspase 3 activation are implicated in neuroprotection by CS-E.     

Development of the auditory sensitivity and formation of the acoustically guided defensive behavior in nestlings of the pied flycatcher Ficedula hypoleuca

Age dynamics of generation of the evoked potentials (EP) in the field L of caudal nidopallium (the higher integrative center of the avian auditory system) and development of the auditory-guided defensive behavior were studied in control and visually deprived pied flycatcher Ficedula hypoleuca nestlings. It was shown that the rhythmically organized monofrequency signals with sound frequency 3.5 kHz and higher produced the defensive behavior as the auditory sensitivity to these frequencies matured. After 9 days, the species-specific alarm signal produced more effectively the defensive behavior than the tonal signals. The rhythmically organized sound with filling frequency 0.5 kHz, occupying the less low-frequency diapason than the feeding signal, produced the effect opposite to the alarm signal to increase the nestling mobility. At the initial stage of the defensive behavior development the auditory threshold fell markedly in the frequency diapason corresponding to the frequency diapason of the alarm signal (5–6 kHz), which seemed to facilitate involvement of this diapason signals in the defensive integration. The auditory EP generation thresholds in the whole studied diapason were lower in the visually deprived nestlings than in the normally developing one; however, the ability of the acoustic signals to suppress alimentary reactions fell significantly.     

Patterns of embryonic neurogenesis in a primitive wingless insect, the silverfish, Ctenolepisma longicaudata: comparison with those seen in flying insects

 Neurogenesis was examined in the central nervous system of embryos of the primitively wingless insect, the silverfish, Ctenolepisma longicaudata, using staining with toluidine blue (TB) and the incorporation of bromodeoxyuridine (BUdR). The silverfish has the same number and positioning of neuroblasts as seen in more advanced insects and the relative order in which the different neuroblasts segregate from the neuroectoderm is highly conserved between Ctenolepisma and the grasshopper, Schistocerca. Of the 31 different neuroblasts found in a thoracic segment, one (NB 6–3) has a much longer proliferative period in silverfish. Of the remainder, 14 have similar proliferative phases, while16 neuroblasts have extended their proliferative period by 10% of embryogenesis or greater in the grasshopper as compared with the silverfish. Both insects had similar periods of abdominal neurogenesis except that in the silverfish terminal ganglion a prominent set of neuroblasts continued dividing until close to hatching, possibly reflecting the importance of cercal sensory input in this insect. This comparison between silverfish and grasshopper shows that the shift from wingless to flying insects was not accompanied by the addition of any new neuronal lineages in the thorax. Instead, selected lineages underwent a proliferative expansion to supply the additional neurons presumably needed for flight. The expansion of specific thoracic lineages was accompanied by the reduction of the terminal abdominal lineages as flying insects began to de-emphasize their cercal sensory system. Received:16 March 1998 / Accepted: 21 May 1998     

Spatially discrete FGF-mediated signalling directs glial morphogenesis.

Neurons provide critical signals that regulate both the number and differentiation of glia. In addition, glia are attracted to and enwrap neuronal axonal processes. FGF-like signalling is thought to be one of the many potential axon-derived morphogenetic signals, however, the multiple roles of FGFs have made experimental tests of these signals difficult in vivo. In the Drosophila FGF receptor mutant heartless, glia migrate to axons, but fail to elongate around them. This study shows that in the similar but larger grasshopper CNS, FGF signalling is likely to mediate one step in the close interaction between glia and axons. FGF2-coated beads attract glia in the CNS and compete with axons for their resident, enwrapped glia. In addition, bath applied FGF2 causes mature axonal glia, which normally enwrap axon tracts, to round up. FGF2 activates the product of the grasshopper heartless FGF receptor gene and probably interferes with the normal function of an endogenous axon-associated FGF-like molecule. It is proposed that insect axons provide a critical spatially restricted FGF-like signal that induces glia to enwrap them.     

听觉中枢神经元对声信号的识别和处理

人类听觉的基本特性和机制与其他哺乳动物相似,因此,利用动物所作的听觉研究和获得的结果,有助于认识人类自身的听觉.围绕听觉中枢神经元对不同模式的声信号的识别和处理,简要综述了这方面的研究.声信号和声模式识别在听觉中枢对声信号的感受和加工中具有重要意义.听神经元作为声模式识别的结构和功能基础,对不同的声刺激模式产生不同反应,甚至是在同一声刺激模式下,改变其中的某个声参数,神经元的反应也会发生相应改变,而其反应的特性和机制均需要更多研究来解答.另外,声信号作为声信息的载体,不同的声信息寓于不同的声参数和声特征之中,研究发现,听觉中枢神经元存在相应的声信息甄别和选择的神经基础,能对动态变化的声频率、幅度和时程等进行反应和编码,并且,在不同种类动物上获得的研究结果极为相似,表明听觉中枢对不同声信号和声刺激模式的识别、分析和加工,具有共同性和普遍性.     

The coding of airborne-sound and vibration signals in bimodal ventral-cord neurons of the grasshopperTettigonia cantans

Summary In grasshoppers, the auditory and vibrational senses converge on the same ventral-cord neurons. All neurons in the ventral cord that discharge impulses in response to either airborne-sound or vibration stimuli also receive synaptic inputs from the other sensory system. The latter elicit either subthreshold excitation or inhibition.The coding of the conspecific song in the responses of most ventral-cord neurons ofTettigonia cantans is considerably improved when the stimulus consists not of simulated natural sounds alone, but of such sounds together with either maintained vibration or vibration matched to the temporal structure of the song.Stridulating tettigoniids produce both airborne and substrate-conducted sound. Thus the perception of airborne sound and vibration, and their simultaneous processing in individual ventral-cord neurons, may be of fundamental importance — not only in localizing a nearby sound source, but also in facilitating the recognition of conspecific signals.Abbreviations SNS simulated natural song The work was supported by the Deutsche Forschungsgemeinschaft, as part of the program Neurale Mechanismen des Verhaltens (Ka 498/1)We are grateful to Dr. M.A. Biederman-Thorson for the translation of the text and to Mrs. A. Scheiding for technical help.     

Thermostimulation-Induced Modulation of Conditioned Reflex Movement-Related Reactions of Cortical Neurons of the Cat

In awake cats trained to perform a food-procuring conditioned operant reflex (placing movement), we studied impulse reactions of 86 neurons of the motor cortex (field 4) related to realization of the above movements. As conditioning stimuli (CS) initiating the reflex, we used either non-noxious electrocutaneous stimulation (ECS) of the contralateral forelimb or an acoustic stimulus (sound click). Impulsation of cortical neurons was recorded under conditions of (i) isolated presentation of the CS (control), (ii) presentation of the CS (either ECS or acoustic stimulus) combined with thermostimulation (heating with a miniature electric bulb) of the skin of the working forelimb, and (iii) the same, but with stimulation of the resting forelimb. When we recorded spike activity of neurons within the projection motor zone of the resting limb subjected to ESC, alternating thermostimulation of both forelimbs resulted in considerable intensification and an increase in the duration of neuronal responses, especially in cases where thermostimulation was applied to the working limb ipsilateral to the recording site (a two- to threefold increase). When spike reactions were recorded within the motor cortex of the working forelimb, thermostimulation resulted in a considerable increase in the intensity of these reactions and a decrease in their latency, but only when such stimulation was applied to the working forelimb. Thermostimulation of the resting (ipsilateral, subjected to ESC) limb evoked opposite effects (the intensity of neuronal reactions dropped). In both situations, placing movements remained within the control limits. When sound click was used as a distant CS, thermostimulation of the working limb enhanced neuronal responses, increased their duration by 50-100%, and also increased the time of forestalling of the movement initiation by spike neuronal reactions. Thermostimulation of the resting forelimb in this situation also suppressed neuronal reactions. We conclude that foreign stimulations directed toward modifications of the receptor model of the operant reflex experimental situation formed in the animal result in a decrease in the intensity of the spike responses of field-4 neurons and prolongation of the latencies of these responses, while stimulations promoting the inflow of afferent information to the cortical projection of the working limb evoke opposite effects, an increase in the intensity of neuronal spike responses and a decrease in their latencies.     

Species Recognition Influences Female Mate Preferences in the Common European Grasshopper (Chorthippus biguttulus Linnaeus, 1758)

Species recognition and intraspecific mating preferences constitute two basic aspects of animal communication. Both can be considered as variations in response to signals and it has been suggested that they represent a continuum. Selection on species recognition could therefore influence intraspecific mating preferences. We show that females of the common European grasshopper Chorthippus biguttulus prefer conspecific male signals that can be distinguished more reliably from sympatrically occurring heterospecific signals. This suggests that in C. biguttulus, sexual selection might be influenced by pleiotropic effects from species recognition. The results show how the heterospecific signal environment could have determined why and in which direction specific traits become sexually selected.     

Substrate-borne Vibrations as a Component of Intraspecific Communication in the Groundhopper <Emphasis Type="Italic">Tetrix ceperoi</Emphasis>

Substrate-borne vibrational signals used in communication of Tetrix ceperoi (Orthoptera: Tetrigidae) are described for the first time. Among all studied Tetrix species, T. ceperoi use the richest repertoire of vibrational signals. These signals consist of isolated pulses or pulses united into rhythmically repeated or irregularly repeated groups and are used in rivalry, pre-copulation, and copulation behaviour (1st, 3rd, and 4th structural types of signals), and are also used by single males or females (2nd type of signal). A 5th type of vibrational signal is produced by wing tremulation, which was unknown Tetrigidae. Based on the results of this study, the vibrational signalisation seems to play an important role in species-specific mate recognition of T. ceperoi. The significance of individual signals is discussed based on results of manipulative contact experiments.     

Role of Acetylcholine in the Modulation of Activity of Neocortical Neurons in Awake Animals Performing an Instrumental Conditioned Reflex

We studied modulatory effects of the cholinergic system on the activity of sensorimotor cortex neurons related to realization of an instrumental conditioned placing reflex. Experiments were carried out on awake cats; multibarrel glass microelectrodes were used for extracellular recording of impulse activity of neurons in the sensorimotor cortex and iontophoretic application of synaptically active agents within the recording region. The background and reflex-related activity was recorded in the course of realization of conditioned movements, and then changes of spiking induced by applications of the testing substances were examined. Applications of acetylcholine and carbachol resulted in increases in the intensity of impulse reactions of neocortical neurons evoked by presentation of an acoustic signal and in simultaneous shortening of the response latencies. An agonist of muscarinic receptors, pylocarpine, exerted a similar effect on the evoked activity of sensorimotor cortex neurons. Blockers of muscarinic receptors, atropine and scopolamine, vice versa, sharply suppressed impulse reactions of cortical neurons to afferent stimulation and simultaneously increased latencies of these responses. Applications of an agonist of nicotinic receptors, nicotine, was accompanied by suppression of impulse neuronal responses, an increase in the latency of spike reactions to presentation of a sound signal, and a corresponding increase in the latency of a conditioned motor reaction. In contrast, application of an antagonist of nicotinic receptors, tubocurarine, significantly intensified neuronal spike responses and shortened their latency. The mechanisms underlying the effects of antagonists of membrane muscarinic and nicotinic cholinoreceptors and the role of activation of these receptors in the modulation of activity of pyramidal and non-pyramidal neocortical neurons related to realization of the instrumental motor reflex are discussed.     

普氏蹄蝠下丘单反应和双反应神经元加工多普勒频移补偿信号的特点及生理机制

恒频-调频(constant frequency-frequency modulation,CF-FM)蝙蝠独特的多普勒频移补偿(Doppler-shift compensation,DSC)行为可保证其对回声信息的精确提取.那么听中枢加工DSC信号的适应性机制是什么?本实验模拟CF-FM蝙蝠DSC后的回声定位信号,研究下丘(inferior colliculus,IC)神经元加工DSC信号的特点及生理机制.实验共获得117个IC神经元,在CF-FM声刺激下,神经元表现为single-on(SO,n=83)和double-on(DO,n=34)两种反应模式.无论是在蝙蝠的正向还是负向补偿过程中,SO和DO神经元对回声反应恢复到50%时的双声刺激间隔(inter-pulse interval,IPI)值,均会随补偿条件的改变而发生变化.当双声刺激由无补偿转变为最佳补偿条件时,两类神经元的50%IPI显著缩短(P0.001),但SO神经元50%IPI缩短率超过70%的神经元数目较DO神经元多,且偏好正向补偿的IC神经元中,SO神经元的平均DSC范围也要显著宽于DO神经元(P0.05).该研究结果提示,IC中SO神经元可能较DO神经元更能充分利用蝙蝠DSC行为,来提高对回声反应的恢复能力,以最大程度地获取猎物信息并准确判断与猎物的相对速度.     

Chondroitin sulfate-D promotes neurite outgrowth by acting as an extracellular ligand for neuronal integrin αVβ3

BackgroundChondroitin sulfate (CS) chains are prominent extra/pericellular matrix components in the central nervous system (CNS) and can exert positive or negative regulatory effects on neurite outgrowth, depending on the CS structure and the amount. Despite the remarkable abilities of highly sulfated forms of CS chains to enhance neurite outgrowth, the neuronal recognition systems for such promotional CS chains, including CS-D polysaccharide, remain to be fully elucidated.MethodsWe explored the molecular basis of the CS-D-mediated neurite extension using primary hippocampal neurons cultured on substrate precoated with CS-D polysaccharides, and evaluated functional involvement of a distinct integrin heterodimer as a novel neuronal CS receptor for CS-D.ResultsWe identified an extracellular matrix receptor, integrin αVβ3, as a functional receptor for CS-D. CS-D, but not CS-C (a precursor form of CS-D) showed significant binding affinity toward recombinant integrin αVβ3 heterodimer and activated intracellular signaling(s) involving focal adhesion kinase (FAK) and Src/Fyn kinase. Functional blockade of the respective players for integrin signaling abrogated the promotional effects of CS-D. We also found the existence of CS-D-induced integrin activation system in neuronal stem/progenitor cell population.ConclusionsThe neuronal cell surface integrin αVβ3 can function as a CS receptor for a highly sulfated CS subtype, CS-D.General significanceOur findings are the first to demonstrate that CS-dependent neurite outgrowth promotion is exerted via direct activation of specific integrin heterodimers on neuronal cell surfaces, providing new insights into understanding the CS-sensing machineries that regulate CNS development and regeneration.     

Neuronal determination during embryonic development of the grasshopper nervous system

We have examined the roles of cell lineage and interactions in the determination of individual identified neurons in the grasshopper embryo by selective ablations of individual cells and/or their neighbors at successive stages following their birth. The neurons in the grasshopper central nervous system (CNS) are produced by two types of identifiable neuronal precursor cells: neuroblasts (NBs), which generate most of the neurons, and midline precursors (MPs), which generate only a few. NBs divide asymmetrically in a stem cell fashion to generate a chain of ganglion mother cells (GMCs) which then divide once more symmetrically to produce pairs of sibling neurons: MPs cleave once to generate a single pair of sibling neurons. We analyzed the determination of (1) the pair of sibling progeny produced by midline precursor 3 (MP3) and the determination of (2) the pair of sibling progeny produced by the first GMC from neuroblast 1-1 (NB 1-1); in each case the siblings normally differentiate into morphologically distinct neurons. Our results indicate that both pairs of neuronal progeny (1) are born equivalent, (2) become determined by cell interactions early in their development before axonogenesis, and (3) demonstrate a hierarchy of fates with one fate dominant over the other. These results suggest a common pattern of neuronal determination in the grasshopper and possibly all insect embryos.     

Pulse-rate recognition in an insect: evidence of a role for oscillatory neurons

Various mechanisms have been proposed as the neural basis for pulse-rate recognition in insects and anurans, including models employing high- and low-pass filters, autocorrelation, and neural resonance. We used the katydid Tettigonia cantans to test these models by measuring female responsiveness on a walking compensator to stimuli varying in temporal pattern. Each model predicts secondary responses to certain stimuli other than the standard conspecific pulse rate. Females responded strongly to stimuli with a pulse-rate equal to half the standard rate, but not to stimuli with double the standard rate. When every second pulse or interval was varied in length, females responded only when the resulting stimuli were rhythmic with respect to the period of the standard signal. These results provide evidence rejecting the use of either high-/low-pass filter networks or autocorrelation mechanisms. We suggest that rate recognition in this species relies on the resonant properties of neurons involved in signal recognition. According to this model, signals with a pulse rate equal to the resonant frequency of the neurons stimulate the female to respond. The results are discussed with regard to both neural and evolutionary implications of resonance as a mechanism for signal recognition. In memory of Dagmar von Helversen (1944–2003)     

Interaction of influences from the auditory and somatosensory afferent systems on neurons of the primary auditory cortex

In experiments on anesthetized cats, 80 neurons of the primary auditory cortex (A1) were studied. Within the examined neuronal population, 66 cells (or 82.5%) were monosensory units, i.e., they responded only to acoustic stimulations (sound clicks and tones); 8 (10.1%) neurons responded to acoustic stimulation and electrocutaneous stimulation (ECS); the rest of the units (7.4%) were either trisensory (responded also to visual stimulation) or responded only to non-acoustic stimulations. In the A1 area, neurons responding to ECS with rather short latencies (15.6–17.0 msec) were found. ECS usually suppressed the impulse neuronal responses evoked by sound clicks. It is concluded that somatosensory afferent signals cause predominantly an inhibitory effect on transmission of an acoustic afferent volley to the auditory cortex at a subcortical level; however, rare cases of excitatory convergence of acoustic and somatosensory inputs toA1 neurons were observed.     

Courtship behavior,communicative sound production and resistance to stress in Drosophila mutants with defective <Emphasis Type="Italic">agnostic</Emphasis> gene coding for LIMK1

To elucidate the role of one of the main elements of signal cascade of actin remodeling—LIM-kinase 1 (LIMK1)—in the control of animal behavior we have studied the characteristics of courtship behavior, parameters of acoustic communicative signals and their resistance to heat shock (HS, 37°C, 30 min) in Drosophila melanogaster males with the agn ^ts3 mutation in the agnostic locus which contains CG1848 gene for LIMK1. The data obtained were compared with the results of our previous similar investigation on wild type CS males [1]. Flies were divided into 4 groups. The males of control groups were not subjected to heat shock. Other groups were comprised of males subjected to heat shock either at the beginning of larval development when the mushroom body neuroblasts are predominantly dividing (groups HS1), or at the prepupal stage when the brain central complex is developing (groups HS2), or at the imago stage 1 hour before the test (groups HS). All males were tested at the age of 5 days. Virgin and fertilized CS females were used as courtship objects. Comparison of control groups of the wild type (CS) and mutant strains has shown that the mutation agn ^ts3 does not affect the main parameters of courtship behavior (courtship latency, the rapidity of achieving copulation, courtship efficiency) but leads to: a decrease in sexual activity, an increase in duration of sound trains in the songs and to a slight increase in the rate and stability of singing pacemakers. When compared to wild type, the agn ^ts3 males are more resistant to HS given 1 hour before the test. After HS their courtship intensity does not decrease and the main parameters of their courtship behavior and communicative sound signals either do not change, or appear to be even better stabilized. The frequency of distorted sound pulses (an indicator of frequency of impairments in the activity pattern of neuro-motor circuits of the singing center) decreases, and the working rate and stability of pacemakers of the pulse and the sine songs increases. Therefore, although the sharply elevated LIMK1 and p-cofilin levels detected by immunofluorescent techniques in the cells of agn ^ts3 mutants can lead to dramatic defects in learning and memory [2] accompanied by a decline of motivation of males, they do not appreciably affect the neuro-motor coordination during singing. The higher resistance of characteristics of the mutant behavior and communicative sound signals to heat shock is in agreement with the fact that the extremely high LIMK1 and P-cofilin levels fall down to normal values after HS and both the learning acquisition and memory formation are restored.     

Corticofugal regulation of auditory sensitivity in the bat inferior colliculus

Under free-field stimulation conditions, corticofugal regulation of auditory sensitivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus, was studied by blocking activities of auditory cortical neurons with Lidocaine or by electrical stimulation in auditory cortical neuron recording sites. The corticocollicular pathway regulated the number of impulses, the auditory spatial response areas and the frequency-tuning curves of inferior colliculus neurons through facilitation or inhibition. Corticofugal regulation was most effective at low sound intensity and was dependent upon the time interval between acoustic and electrical stimuli. At optimal interstimulus intervals, inferior colliculus neurons had the smallest number of impulses and the longest response latency during corticofugal inhibition. The opposite effects were observed during corticofugal facilitation. Corticofugal inhibitory latency was longer than corticofugal facilitatory latency. Iontophoretic application of γ-aminobutyric acid and bicuculline to inferior colliculus recording sites produced effects similar to what were observed during corticofugal inhibition and facilitation. We suggest that corticofugal regulation of central auditory sensitivity can provide an animal with a mechanism to regulate acoustic signal processing in the ascending auditory pathway. Accepted: 15 July 1998