Descending brain neurons in the cricket Gryllus bimaculatus (de Geer): auditory responses and impact on walking

The activity of four types of sound-sensitive descending brain neurons in the cricket Gryllus bimaculatus was recorded intracellularly while animals were standing or walking on an open-loop trackball system. In a neuron with a contralaterally descending axon, the male calling song elicited responses that copied the pulse pattern of the song during standing and walking. The accuracy of pulse copying increased during walking. Neurons with ipsilaterally descending axons responded weakly to sound only during standing. The responses were mainly to the first pulse of each chirp, whereas the complete pulse pattern of a chirp was not copied. During walking the auditory responses were suppressed in these neurons. The spiking activity of all four neuron types was significantly correlated to forward walking velocity, indicating their relevance for walking. Additionally, injection of depolarizing current elicited walking and/or steering in three of four neuron types described. In none of the neurons was the spiking activity both sufficient and necessary to elicit and maintain walking behaviour. Some neurons showed arborisations in the lateral accessory lobes, pointing to the relevance of this brain region for cricket audition and descending motor control.     

Developmental experience alters information coding in auditory midbrain and forebrain neurons

In songbirds, species identity and developmental experience shape vocal behavior and behavioral responses to vocalizations. The interaction of species identity and developmental experience may also shape the coding properties of sensory neurons. We tested whether responses of auditory midbrain and forebrain neurons to songs differed between species and between groups of conspecific birds with different developmental exposure to song. We also compared responses of individual neurons to conspecific and heterospecific songs. Zebra and Bengalese finches that were raised and tutored by conspecific birds, and zebra finches that were cross‐tutored by Bengalese finches were studied. Single‐unit responses to zebra and Bengalese finch songs were recorded and analyzed by calculating mutual information (MI), response reliability, mean spike rate, fluctuations in time‐varying spike rate, distributions of time‐varying spike rates, and neural discrimination of individual songs. MI quantifies a response's capacity to encode information about a stimulus. In midbrain and forebrain neurons, MI was significantly higher in normal zebra finch neurons than in Bengalese finch and cross‐tutored zebra finch neurons, but not between Bengalese finch and cross‐tutored zebra finch neurons. Information rate differences were largely due to spike rate differences. MI did not differ between responses to conspecific and heterospecific songs. Therefore, neurons from normal zebra finches encoded more information about songs than did neurons from other birds, but conspecific and heterospecific songs were encoded equally. Neural discrimination of songs and MI were highly correlated. Results demonstrate that developmental exposure to vocalizations shapes the information coding properties of songbird auditory neurons. © 2009 Wiley Periodicals, Inc. Develop Neurobiol 70: 235–252, 2010.     

Brain projections and information processing of biologically significant sounds by two large ventral-cord neurons ofGryllus bimaculatus DeGeer (Orthoptera,Gryllidae)

Summary Two ventral-cord neurons in the auditory system ofGryllus bimaculatus were studied electrophysiologically by stimulation with pulses of sound at a single frequency (sine-wave pulses), stridulatory songs, and artificial sounds constructed to imitate the conspecific songs. The sine-wave pulses were varied in frequency, sound intensity, duration, and repetition rate. The stridulatory songs were the conspecific calling, aggressive, and courtship songs and the calling songs of 8 sympatric gryllids (played back at different sound intensities). The artificial songs were varied in carrier frequency, pulse rate, chirp rate, and sound intensity.The LF₁ neuron precisely duplicates the temporal structure of the conspecific calling (and aggressive) song over the whole intensity range (Figs. 7, 8, 10). It is sharply tuned to the carrier frequency of the song (5 kHz) and shows little or no response above 10 kHz and below 3 kHz (Figs. 1, 2). By variation of the calling song's temporal structure it can be demonstrated that the LF₁ neuron is particularly suited to respond to the pulse duration and the pulse and chirp repetition rates of this song pattern (Figs. 6, 9).On the other hand, the HF₁ neuron is a broad-band neuron with a maximal sensitivity at 16 kHz (Figs. 1, 4); it is tuned to the conspecific courtship song with respect to carrier frequency, the short pulse duration, and the very low pulse repetition rate (Figs. 6, 7, 8).The results demonstrate that the two ventral-cord neurons represent highly evolved channels of the auditory pathway in gryllids, each of which transmits important features of the corresponding conspecific songs to several areas of the brain (Fig. 11). But they are not ideal filters for these conspecific songs, since they also respond to many other sound signals (Fig. 10).Supported by the Deutsche Forschungsgemeinschaft as part of the program Sonderforschungsbereich 114 (Bionach), BochumUnder the auspices of the scientist exchange program of the Deutsche Forschungsgemeinschaft and the Academy of Sciences, USSRWe thank Prof. Dr. Schwartzkopff for his help and support; it was due to his initiative and organization that this work could be done in collaboration between the Sechenov Institute, Leningrad, and the Lehrstuhl für Allgemeine Zoologie, Ruhr University, Bochum. We are grateful to Mrs. I. Klotz and Mrs. B. Brücher for technical assistance.     

Sex pheromone and plant-associated odour processing in antennal lobe interneurons of male Spodoptera littoralis (Lepidoptera: Noctuidae)

Antennal lobe interneurons of male Spodoptera littoralis (Boisd.) were investigated by using intracellular recording and staining techniques. Physiological and morphological characteristics of local interneurons and projection neurons responding to sex pheromone and plant-associated volatiles are described. The interneurons identified were divided into three groups, depending on their physiological response characteristics. Both types of interneurons, local interneurons and projection neurons, were described in all three groups. 1. Interneurons responding exclusively to sex pheromone stimuli, displayed different degrees of specificity. These neurons responded to either one, two, three or all four of the single sex pheromone or sex pheromone-like compounds tested. Most of these neurons also responded to a mixture of the two pheromone components present in the female S. littoralis blend. Two local interneurons and one projection neuron were identified as blend specialists, not responding to the single female produced sex pheromone components, but only to their mixture. Five pheromone specific projection neurons arborized in one or more subcompartments of the macroglomercular complex (MGC) and some of them had axonal branches in the calyces of the mushroom body and in different parts of the lateral protocerebrum. 2. Interneurons responding only to plant-associated volatiles varied highly in specificity. Neurons responding to only one of the stimuli, neurons responding to a variety of different odours and one neuron responding to all stimuli tested, were found. Three specialized local interneurons had arborizations only in ordinary glomeruli. One specialized and three less specialized local interneurons had arborizations within the MGC and the ordinary glomeruli. The projection neurons responding only to plant-associated volatiles had mostly uni- or multiglomerular arborizations within the ordinary glomeruli. 3. Interneurons responding to both sex pheromones and plant-associated stimuli varied in specificity. Individual interneurons that responded to few plant-associated odours mostly responded to several pheromone stimuli as well. Projection neurons responding to most of the plant-associated volatiles also responded to all pheromone stimuli. Two local interneurons responding to both stimulus groups, arborized within the MGC and the ordinary glomeruli. Projection neurons mostly arborized in only one ordinary glomerulus or in one compartment of the MGC. The variation in specificity and sensitivity of antennal lobe interneurons and structure-function correlations are discussed.     

Sexual differentiation and seasonal variation in response to conspecific and heterospecific acoustic signals

Interspecific territoriality is frequently reported between closely related species; however, few studies have demonstrated interspecific territoriality between distantly related species living in sympatry. We conducted playback experiments to investigate territorial behaviour in male and female White‐bellied Wrens (Uropsila leucogastra) in response to simulated conspecific and heterospecific intruders during the breeding and non‐breeding seasons. We explored whether heterospecific songs of the Happy Wren (Pheugopedius felix), a distantly related species and ecological competitor, elicited antagonistic responses from focal White‐bellied Wrens, and whether such responses differed between the sexes. We also examined whether male and female responses to conspecific and heterospecific rivals varied with season. We found that male White‐bellied Wrens always responded to conspecific song, and responded significantly more to heterospecific song compared to a control stimulus (Tropical Parula, Setophaga pitiayumi). In contrast, although female White‐bellied Wrens responded strongly to conspecific song, their response to heterospecific song did not differ significantly from the control stimulus. The proportion of males that responded to heterospecific songs and the proportion of females that responded to conspecific songs varied seasonally, showing significantly lower responses during the breeding season. The intense responses of male White‐bellied Wrens to playback of heterospecific songs suggest that they recognise ecological competitors based on their vocal signals. Furthermore, the decrease in agonistic interactions during the breeding season is in line with the hypothesis that aggressive behaviour may be detrimental to reproductive and parental activity, and the hypothesis that heterospecific animals pose less of a threat during the breeding season.     

Temporal processing of vibratory communication signals at the level of ascending interneurons in Nezara viridula (Hemiptera: Pentatomidae)

During mating, males and females of N. viridula (Heteroptera: Pentatomidae) produce sex- and species-specific calling and courtship substrate-borne vibratory signals, grouped into songs. Recognition and localization of these signals are fundamental for successful mating. The recognition is mainly based on the temporal pattern, i.e. the amplitude modulation, while the frequency spectrum of the signals usually only plays a minor role. We examined the temporal selectivity for vibratory signals in four types of ascending vibratory interneurons in N. viridula. Using intracellular recording and labelling technique, we analyzed the neurons' responses to 30 pulse duration/interval duration (PD/ID) combinations. Two response arrays were created for each neuron type, showing the intensity of the responses either as time-averaged spike counts or as peak instantaneous spike rates. The mean spike rate response arrays showed preference of the neurons for short PDs (below 600 ms) and no selectivity towards interval duration; while the peak spike rate response arrays exhibited either short PD/long ID selectivity or no selectivity at all. The long PD/short ID combinations elicited the weakest responses in all neurons tested. No response arrays showed the receiver preference for either constant period or duty cycle. The vibratory song pattern selectivity matched the PD of N. viridula male vibratory signals, thus pointing to temporal filtering for the conspecific vibratory signals already at level of the ascending interneurons. In some neurons the responses elicited by the vibratory stimuli were followed by distinct, regular oscillations of the membrane potential. The distance between the oscillation peaks matched the temporal structure of the male calling song, indicating a possible resonance based mechanism for signal recognition.     

Prothoracic DUM neurons of the cricket Gryllus bimaculatus — responses to natural stimuli and activity in walking behavior

Summary Responses to sensory stimuli and spike activity uring walking were investigated in bilaterally symmetrical dorsal unpaired median (DUM) neurons of the cricket. Intracellular recordings within the prothoracic ganglion were made either in restrained animals or in stationary walking specimens whilst parameters of their intended locomotion were measured. Three types of DUM cells were distinguished morphologically and physiologically. DUMa neurons send axons through segmental nerves. They often generated spontaneously large action potentials with low frequencies. Most DUMa neurons showed multimodal sensitivity, preferentially to cereal wind puffs and 15 kHz sound. Mean latencies ranged from 25 to 349 ms. Their large intraindividual variability could be correlated with behavioral modes during walking. Generally, the spike frequency increased with increased forward speed, while it was not related to turning. DUMb neurons projected either through the anterior or posterior connectives, but seemed physiologically similar to DUMa neurons. DUMc neurons were H-shaped with axons in both pairs of connectives. No external stimulus led to discrete spikes, but the regular spontaneous activity was modulated following cereal wind puffs to a restrained animal. During wind evoked escape the spike activity of another DUMc cell was modulated in phase with the rhythmic running behavior. The possibly different functions of DUMa and DUMc neurons during walking are discussed.     

Effects of self-generated wind on compensational recovery of escape direction in unilaterally cercus-ablated crickets, Gryllus bimaculatus

The effects of self-generated wind on the compensational recovery of escape direction were investigated in unilaterally cercus-ablated crickets, Gryllus bimaculatus. To separate walking and self-generated wind during walking, unilaterally cercus-ablated crickets were placed on a styrofoam ball that was easily rotated by leg motion during walking. The stationary walking on the ball did not produce self-generated wind, because no body motion occurred. Crickets that were trained on the ball but given no artificial air puff for 14 days after cercal ablation did not show any compensational recovery of escape direction. Therefore, spontaneous walking itself was not sufficient to compensate the wind-evoked escape direction in the crickets. Artificial air puffs from the anterior direction synchronized with the stationary walking were effective for the compensational recovery of escape direction, but those from the posterior direction were not. As most of the spontaneous walking was directed to the forward direction, only an artificial air puff from the anterior direction coincided well with actual self-generated wind occurring during the onset of normal walking. Therefore, self-generated wind during walking seems essential for the compensational recovery of escape direction in unilaterally cercus-ablated crickets. When artificial air puffs were unsynchronized with walking, no compensational recovery was observed. This result suggests that artificial air puffs should be given just after the onset of spontaneous walking. Otherwise, the artificial air puffs may not be recognized as self-generated wind.     

Auditory nerve and interneurone responses to natural sounds in several species of cicadas

ABSTRACT. The calling and courtship songs of 17-year cicadas and of Say's cicadas differ both in the sound frequency spectrum and in temporal pattern. Multiunit recordings with hook electrodes from the whole auditory nerve show that the hearing organs are especially sensitive to transient stimuli occurring in natural sounds. Artificially produced clicks elicit bursts of spikes synchronized among various primary sensory fibres. These fibres respond to natural calling and courtship songs with a specificity dependent on carrier frequency, rhythm and transient content of the sound, following sound pulses (i.e. tymbal actions) up to repetition rates of 200 Hz. An ascending, plurisegmental interneurone was characterized by intracellular recording and simultaneously stained with cobalt. Its main arborization spatially overlaps the anterior part of the sensory auditory neuropile, and the axon was traced as far as the prothoracic ganglion. Direct input from primary auditory fibres was suggested by latency measurements. Intracellular recordings from such neurons in different species show distinct auditory input, with phasic-tonic spike responses to tones. In general, the interneurone response is more species-specific to calling than to courtship songs, and the preferential response to the conspecific calling song is based primarily upon sound frequency content.     

The function of auditory neurons in cricket phonotaxis

Summary The activity of auditory receptor cells and prothoracic auditory neurons of the cricket,Gryllus bimaculatus, was recorded intracellularly while the animal walked on a sphere or while passive movement was imposed on a foreleg.During walking the responses to simulated calling song is altered since (i) the auditory sensory cells and interneurons discharged impulses in the absence of sound stimuli (Figs. 1, 3) and (ii) the number of action potentials in response to sound is reduced in interneurons (Figs. 2, 3).These two effects occurred in different phases of the leg movement during walking and therefore masked, suppressed or did not affect the responses to auditory stimuli (Figs. 3, 4). Hence there is a time window within which the calling song can be detected during walking (Fig. 5).The extra excitation of receptors and interneurons is probably produced by vibration of the tympanum because (i) the excitation occurred at the same time as the leg placement (Fig. 4), (ii) during walking on only middle and hindlegs, no extra action potentials were observed (Fig. 6), (iii) in certain phases of passive movements receptor cells and interneurons were excited as long as the ipsilateral ear was not blocked (Figs. 8, 9).Suppression of auditory responses seems to be peripheral as well as central in origin because (i) it occurred at particular phases during active and passive leg movements in receptor cells and interneurons (Figs. 1, 4, 9), (ii) it disappeared if the ear was blocked during passive leg movements (Fig. 9) and (iii) it persisted if the animal walked only on the middle and hind legs (Fig. 6).     

Species-Recognition in the Field Cricket, Teleogryllus oceanicus: Behavioral and Neural Mechanisms

SYNOPSIS. Field crickets depend on acoustic organs to detectthe presence of potential predators as well as conspecific crickets.Predators are recognized largely on the basis of spectral frequenciesthat are contained in their acoustic signals. Puffs of air andvery low frequencies activate a cricket's cereal receptors andultrasonic frequencies activate their tympanal organs. Bothof these acoustic stimuli release "escape behavior," in theform of evasive movements. An identified neuron sensitive toultrasound is described. Crickets recognize singing conspecificsby both frequency and temporal properties of cricket songs;however species recognition requires specific temporal informationin calling songs. While previous studies have emphasized therole of songs on female behavior, males also recognize conspecificsongs; sexual differences in recognition behavior occur.     

Species Recognition During Substrate-Borne Communication in <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.) (Pentatomidae: Heteroptera)

The information code in the temporal and spectral characteristics of the substrate-borne communication signals produced by insects has been primarily studied in insects in the suborder Auchenorrhyncha. In the present study we investigated which of the female calling song (FCS) parameters in Nezara viridula (L.) (Heteroptera, Pentatomidae) are essential for recognition by conspecific males. In playback experiments we measured male vibrational responsiveness to FCS signals varying in the durations of pulse trains and inter-pulse train intervals, repetition times, duty cycles, and dominant frequencies, and determined the preference range for each specific parameter. Males were able to distinguish songs of different temporal and frequency parameters and responded best to values characteristic of the song of conspecific females. Signal recognition is achieved on the basis of two temporal filters tuned to the durations of the pulse train and inter-pulse train interval. Males responded best to the dominant frequency characteristic of conspecific songs, which are tuned to the resonant properties of the herbaceous plants used for intraspecific signal transmission during communication.     

Song discrimination learning in zebra finches induces highly divergent responses to novel songs

Perceptual biases can shape the evolution of signal form. Understanding the origin and direction of such biases is therefore crucial for understanding signal evolution. Many animals learn about species-specific signals. Discrimination learning using simple stimuli varying in one dimension (e.g. amplitude, wavelength) can result in perceptual biases with preferences for specific novel stimuli, depending on the stimulus dimensions. We examine how this translates to discrimination learning involving complex communication signals; birdsongs. Zebra finches (Taeniopygia guttata) were trained to discriminate between two artificial songs, using a Go/No-Go procedure. The training songs in experiment 1 differed in the number of repeats of a particular element. The songs in experiment 2 differed in the position of an odd element in a series of repeated elements. We examined generalization patterns by presenting novel songs with more or fewer repeated elements (experiment 1), or with the odd element earlier or later in the repeated element sequence (experiment 2). Control birds were trained with only one song. The generalization curves obtained from (i) control birds, (ii) experimental birds in experiment 1, and (iii) experimental birds in experiment 2 showed large and systematic differences from each other. Birds in experiment 1, but not 2, responded more strongly to specific novel songs than to training songs, showing 'peak shift'. The outcome indicates that learning about communication signals may give rise to perceptual biases that may drive signal evolution.     

蜜蜂下行神经元的光反应特征

利用电生理细胞外记录方法研究了蜜蜂下行神经元对光的反应特征以及与水平方向上光刺激角度的关系。 光刺激主要引起下行神经元的给光反应和撤光反应,持续性反应很弱。下行神经元的反应依赖于光刺激强度,在一定的强度范围内,随着光强度增大,反应强度也增大根据改变刺激角度引起的反应,在水平方向上最敏感,张角最大,约大于40°。     

Neural Basis of Stimulus-Angle-Dependent Motor Control of Wind-Elicited Walking Behavior in the Cricket Gryllus bimaculatus

Crickets exhibit oriented walking behavior in response to air-current stimuli. Because crickets move in the opposite direction from the stimulus source, this behavior is considered to represent ‘escape behavior’ from an approaching predator. However, details of the stimulus-angle-dependent control of locomotion during the immediate phase, and the neural basis underlying the directional motor control of this behavior remain unclear. In this study, we used a spherical-treadmill system to measure locomotory parameters including trajectory, turn angle and velocity during the immediate phase of responses to air-puff stimuli applied from various angles. Both walking direction and turn angle were correlated with stimulus angle, but their relationships followed different rules. A shorter stimulus also induced directionally-controlled walking, but reduced the yaw rotation in stimulus-angle-dependent turning. These results suggest that neural control of the turn angle requires different sensory information than that required for oriented walking. Hemi-severance of the ventral nerve cords containing descending axons from the cephalic to the prothoracic ganglion abolished stimulus-angle-dependent control, indicating that this control required descending signals from the brain. Furthermore, we selectively ablated identified ascending giant interneurons (GIs) in vivo to examine their functional roles in wind-elicited walking. Ablation of GI8-1 diminished control of the turn angle and decreased walking distance in the initial response. Meanwhile, GI9-1b ablation had no discernible effect on stimulus-angle-dependent control or walking distance, but delayed the reaction time. These results suggest that the ascending signals conveyed by GI8-1 are required for turn-angle control and maintenance of walking behavior, and that GI9-1b is responsible for rapid initiation of walking. It is possible that individual types of GIs separately supply the sensory signals required to control wind-elicited walking.     

Asymmetric song responses in coexisting Japanese cicadas

Diel partitioning of singing, which has been observed from the tropics to the temperate regions, is among the various mechanisms of sound partitioning found in multispecies cicada assemblages and is regarded as an important mechanism of coexistence. Using playback experiments, we studied interspecific interference between two Japanese cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata, which exhibit similar peak frequencies in their calling songs and different diel patterns of song activity. We found that the males of the two species had different song response patterns: C. facialis responded significantly more to conspecific calling songs than G. nigrofuscata. These results suggest that male C. facialis have evolved more intense conspecific song recognition than G. nigrofuscata. This may cause male G. nigrofuscata to avoid destructive acoustic interference during their active singing time.     

The effect of ultrasound on the attractiveness of acoustic mating signals

Abstract. Previous laboratory studies ( Nolen & Hoy 1986b ) have shown that the phonotactic responses of flying crickets are influenced by the relative intensities of attractive (mating signal) and repulsive (predator) stimuli. At the functional level, these results suggest that predator cues (ultrasound) can change the attractiveness of a calling song. Using extracellular recordings from cervical connectives it was shown that, like other field crickets, Gryllus rubens (south-eastern field cricket) is sensitive to ultrasound. This ultrasonic sensitivity has probably evolved in response to predation pressure from echolocating bats. Using acoustic playback under field conditions, it was tested whether the relative attractiveness of two male calling songs was influenced by the simultaneous broadcast of ultrasound. A simulated male calling song of G. rubens was broadcast at two different intensities (109 and 103 dB) from two sound traps that caught flying crickets attracted to the songs. Simulated bat ultrasound was broadcast simultaneously with the high-intensity calling song (109 dB) and the relative catch in each of the two traps was measured. The intensity of the ultrasound was varied on different nights. The relative attractiveness of the high-intensity sound trap decreased significantly as the intensity of the ultrasound broadcast with it was increased. For the lowest of the ultrasound broadcast levels, the relative attractiveness did not differ from that expected for two calling songs broadcast without ultrasound. Thus, increased levels of simulated predation risk decreased the attractiveness of the calling song associated with it. These are the first field experiments to show that predation risk in the form of simulated bat ultrasound influences the phonotactic behaviour of flying crickets.     

Interactions of mechanical stimuli and sex pheromone information in antennal lobe neurons of a male moth,<Emphasis Type="Italic"> Spodoptera littoralis</Emphasis>

Male moths respond to sex pheromone sources with up-wind flight behaviour. Localization of the odour source requires not only detection of the olfactory stimulus, but also other sensory input regarding, e.g. visual and mechanical stimuli. Thus, integration of different types of sensory input is necessary. It is, however, not known where in the central nervous system the integration of information regarding different sensory modalities takes place. Using intracellular recording and staining techniques, we investigated neurons in the antennal lobe of Spodoptera littoralis, during stimulation with a mechanical stimulus and a sex pheromone. Fifteen percent of all the neurons investigated responded to the mechanical stimulus and the majority of these neurons showed altered responses if the olfactory stimulus was added. A receptor neuron responding only to the wind stimulus was found to arborise in the antennal lobe. Most projection neurons responded with an enhanced action potential frequency to the combined stimulus. In local interneurons, enhancement, depression, or no change of the responses to the wind stimulus was found when the olfactory stimulus was added. The results suggest that neurons present in the antennal lobe integrate mechanosensory and olfactory input, possibly assisting the moths to orient during up-wind flight towards an odour source.     

Processing of Song Signals in the Cricket and its Hormonal Control

SYNOPSIS. Phonotaxis by female crickets to the calling songof males, is an important model for investigating the neuralbasis of auditory behavior. Recent advances make it possibleto explain some components of this behavior and its hormonalcontrol, at the level of identified neurons and molecular expressionwithin those neurons Tonotopically arranged afferents from the cricket's ear, projectto local and intersegmental prothoracic interneurons. Bilateralprocessing of signals and some temporal-pattern specific processingoccurs in the prothoracic ganglion and influences acoustic informationthat is sent to the brain via ascending interneurons that aredemonstrably involved in phonotaxis. High, low and band- passinterneurons in the brain continue temporal pattern processingwhich matches the selectivity of phonotaxis and may be filtersfor recognition of the calling song. Neurons descending fromthe brain and prothoracic ganglion, direct multimodal signals(including auditory) to more posterior regions, possibly theleg motor neurons that are responsible for phonotaxis Age-related changes or artificially induced changes in JuvenileHormone III levels regulate the threshold for phonotaxis inAcheta domesticus, by varying the threshold of LI, a prothoracicascending interneuron that is necessary for phonotaxis to lowintensity calling songs. Results from in situ hybridizationsuggest that this might be accomplished, in part, by controllingthe levels of nicotinic acetylcholine receptor-like mRNA expressedin LI, presumably by increasing its neurotransmitter receptordensity. L3 is a prothoracic ascending interneuron that exhibitsbandselective response properties to the syllable period ofthe calling song. L3's response is age and JHIII related, andis correlated to phonotactic selectivity. These changes in L3might be accomplished, at least in part by JHIII regulatingthe expression of nicotinic acetylcholine receptor-like mRNAin L3     

Dose-dependent response characteristics of antennal lobe neurons in the male moth Agrotis segetum (Lepidoptera: Noctuidae)

Responses of antennal lobe neurons to different amounts of the female-produced pheromone blend and to its individual components were investigated in Agrotis segetum males using intracellular recording methods. We identified three physiological types of antennal lobe neurons, categorized according to their response thresholds to single pheromone components and to the blend: generalist neurons, component-specific neurons and blend-specific neurons. Response and specificity of antennal lobe neurons were largely dose dependent. In most cases specific responses occurred only at low stimulus amounts, while increasing concentrations often resulted in an increase of the number of pheromone stimuli to which the neuron responded. Dose-response relationships often differed between different stimuli activating a neuron. Accepted: 24 May 1997