The jamming avoidance response in gymnotoid pulse-species: A mechanism to minimize the probability of pulse-train coincidence

Summary Gymnotoid electric fish with pulse-type electric organ discharges (EODs) shorten (lengthen) their EOD intervals as pulses of a slightly slower (faster) train scan their EODs (Figs. 1, 2). They thus minimize the chance of pulse coincidence by transient accelerations (decelerations) of their EOD rate.Studies in curarized preparations demonstrate that this Jamming Avoidance Response (JAR) is controlled by electroreceptive input alone and without reference to an internal electric organ pacemaker-related signal (Fig. 8). A sufficient stimulus input consists of a train of strong, EOD-like stimulus pulses (S₁), which mimic the animal's experience of its own EOD, and a train of small pulses (S₂) of slightly different repetition rate, which mimic EODs of a neighbor. Correct behavioral responses require S₁ pulses of sufficient intensity to recruit pulse-markertype receptors; also spatial and temporal patterns must closely resemble those of the animal's EOD. These features are of little significance for S₂ pulses which, while scanning S₁ pulses, only provide a small perturbation of electroreceptive feedback from S₁ pulses. Inappropriate S₁ stimulation impairs and sometimes reverses (Fig. 7) the behavioral discrimination of scan directions. The JAR is explained in terms of excitatory and inhibitory processes (Fig. 3) which are triggered by S₂ stimulation, at specific phases within the S₁ cycle (Figs. 4–6).The JAR in pulse species strongly resembles the JAR in wave-species (Bullock et al., 1972) and could be considered an evolutionary ancestor of the latter. It is a response to a particular novelty in electroreceptive feedback.We thank Drs. T.H. Bullock, C. Hopkins and an anonymous referee for most helpful criticism. This research was supported by NSF grand BMS74-18640 and NIMH grant PHSMH-2614901 to W.H. and NIH grant/ROI NS 12337-01 to J.B.     

Sensory cues for the gradual frequency fall responses of the gymnotiform electric fish, Rhamphichthys rostratus

The sensory cues for a less known form of frequency shifting behavior, gradual frequency falls, of electric organ discharges (EODs) in a pulse-type gymnotiform electric fish, Rhamphichthys rostratus, were identified. We found that the gradual frequency fall occurs independently of more commonly observed momentary phase shifting behavior, and is due to perturbation of sensory feedback of the fish's own EODs by EODs of neighboring fish. The following components were identified as essential features in the signal mixture of the fish's own and the neighbor's EOD pulses: (1) the neighbor's pulses must be placed within a few millisecond of the fish's own pulses, (2) the neighbor's pulses, presented singly at low frequencies (0.2–4 Hz), were sufficient, (3) the frequency of individual pulse presentation must be below 4 Hz, (4) amplitude modulation of the sensory feedback of the fish's own pulses induced by such insertions of the neighbor's pulses must contain a high frequency component: sinusoidal amplitude modulation of the fish's own EOD feedback at these low frequencies does not induce gradual frequency falls. Differential stimulation across body surfaces, which is required for the jamming avoidance response (JAR) of wave-type gymnotiform electric fish, was not necessary for this behavior. We propose a cascade of high-pass and low-pass frequency filters within the amplitude processing pathway in the central nervous system as the mechanism of the gradual frequency fall response.Abbreviations EOD electric organ discharge - f frequency of EOD or pacemaker command signal - JAR jamming avoidance response - S ₁ stimulus mimicking fish's own EOD - f ₁ frequency of S₁ - S ₂ stimulus mimicking neighbor's EOD - f ₂ frequency of S₂     

‘Recognition units’ at the top of a neuronal hierarchy?

The electric fish, Eigenmannia, is able to discriminate the sign of the frequency difference, Df, between a neighbor's electric organ discharges (EODs) and its own. The fish lowers its EOD frequency for positive Dfs and raises its frequency for negative Dfs to minimize jamming of its electrolocation ability by a neighbor's EODs of similar frequency. This jamming avoidance response (JAR) is controlled by a group of 'sign-selective' neurons in the prepacemaker nucleus (PPN) that is located at the boundary of the midbrain and the diencephalon (Fig. 1). Extracellular recordings from a total of 35 neurons revealed a great similarity between behavioral and neuronal response properties: 1. All neurons fired vigorously for negative Dfs and were almost silent for positive Dfs, regardless of the orientation of the jamming stimulus, and thus discriminated the sign of Df unambiguously (Fig. 2). 2. In accordance with behavioral observations, individual neurons failed to discriminate the sign of Df when the jamming stimulus had the same field geometry as the signal mimicking the animal's own EOD (Fig. 3). 3. Df magnitudes which evoke strongest JARs, usually 4 to 8 Hz, also induced most vigorous responses in sign-selective neurons (Fig. 5). 4. Behavioral and neuronal thresholds for the detection of small jamming signals were similar. Threshold for sign selectivity was reached when the amplitude ratio of the jamming signal to the EOD mimic, measured near the head surface, was 0.001. This value corresponds to a maximal temporal disparity (a necessary cue for performing a correct JAR) of 1 to 2 microseconds for signals received by the two sides of the body in a transverse jamming field (Fig. 7). 5. The effects of two jamming fields, offered orthogonally to each other, may interact nonlinearly at the behavioral as well as at the neuronal level. A positive Df presented in one field may suppress behavioral and neuronal responses to modulations of the sign of Df in the other field (Fig. 8c).     

Electrocommunication and Social Behaviour in Marcusenius senegalensis (Mormyridae,Teleostei)

The electric organ discharges (EODs) of Marcusenius senegalensis, a West African freshwater fish, are bipolar pulses of short duration (220 ± SE 13 μs). In males (n = 10; 10.1–13.1 cm standard length — which is around the size of getting mature), the duration of EOD pulses was of significantly greater variance than in females (n = 9; 9.8–12.8 cm standard length). Male EODs also showed a tendency for a longer duration than female EODs. Groups of three as well as of 14 M. senegalensis formed temporary schools in a ‘naturally’ equipped 720-1 tank. While swimming slowly in a loose school during their nocturnal active phase, fish discharged in irregular long-short-long inter-EOD interval patterns. Near neighbours displayed a tendency to discharge in intervals of similar duration (nearest neighbour distance < 1/2 fish length). On removal of a plastic partition that had separated a pair of fish for at least 3 days, mutual threat displays followed by fighting were observed. During threatening, the fish alternated regularly between bursts of a high discharge rate and short discharge breaks; the rate of change was 4/s. The subdominant animal in a group of two was attacked frequently and often ceased discharging when the dominant fish approached. Courtship behaviour involving gonadally mature fish was accompanied by high-discharge-rate displays with intervals of constant duration in both fish, and the reciprocal display of ‘preferred’ EOD latencies in the 12 ms range. The results demonstrate electric communication by distinct inter-discharge interval patterns in the social behaviour of this mormyrid fish.     

Trained Weakly-electric Fishes Pollimyrus isidori and Gnathonemus petersii (Mormyridae,Teleostei) Discriminate between Waveforms of Electric Pulse Discharges

Fish of the family Mormyridae emit weak, pulse-like electric organ discharges (EODs). The discharge rhythm is variable, but the waveform of the EOD is constant for each fish, with species- and individual characteristics. The ability of Pollimyrus isidori and Gnathonemus petersii (Mormyridae) to discriminate between different EOD waveforms was tested using a differential conditioning procedure. Fish were first trained to respond to a reference signal in swimming to a dish to receive a bloodworm (food reward). The reference signal consisted of a 10-Hz train of the digitally recorded EOD of a conspecific. Second, an alternative signal (10-Hz train of a different EOD, either from another species, or from a conspecific of the other sex) was associated with air bubbles as punishment. The two signals were played at successive trials in random order. On each trial the latency was measured between the onset of the signal and the response. 7 out of the 8 P. isidori tested and both of the two G. petersii tested associated the reference EOD with food. Among these, five P. isidori and two G. petersii responded differentially (p < 0.01) to EODs of different species. P. isidori similarly discriminated between conspecific EODs of different sexes. The quantity of different alternative EODs which could be tested was limited when fish eventually habituated to the punishment. Even when the amplitude of the EODs was randomly changed at each trial, two out of two G. petersii differentiated between EODs of the two species, and three out of three P. isidori tested differentiated between EODs within their own species. Response latencies to the rewarded signal during the basic training and during discrimination (when it had to be distinguished from the S-) were similar. G. petersii showed differential responses for S+ and S- also in the rhythm of discharge exhibited during playback, after five EOD pulses for one fish, and after a single pulse for the other. Mormyrids may therefore distinguish between conspecifics and members of other species, and even between individual conspecifics, by their EOD waveform.     

Distinct mechanisms of modulation in a neuronal oscillator generate different social signals in the electric fishHypopomus

Summary The medullary pacemaker nucleus of the gymnotiform electric fish,Hypopomus, is a relatively simple neuronal oscillator which contains pacemaker cells and relay cells. The pacemaker cells generate a regular discharge cycle and drive the relay cells which trigger pulse-like electric organ discharges (EODs). The diencephalic prepacemaker nucleus (PPN) projects to the pacemaker nucleus and modulates its activity to generate a variety of specific discharge patterns which serve as communicatory signals (Figs. 2 and 3).While inducing such signals by microiontophoresis of L-glutamate to the region of the PPN (Fig. 4) of curarized animals, we monitored the activity of neurons in the pacemaker nucleus intracellularly. We found that pacemaker cells and relay cells were affected differently in a manner specific to the type of EOD modulation (Figs. 5–10). The normal sequence of pacemaker cell and relay cell firing was maintained during gradual rises and falls in discharge rate. Both types of cells ceased to fire during interruptions following a decline in discharge rate. During sudden interruptions, however, relay cells were steadily depolarized, while pacemaker cells continued to fire regularly. Short and rapid barrages of EODs, called chirps, were generated through direct and synchronous activation of the relay cells whose action potentials invaded pacemaker cells antidromically and interfered with their otherwise regular firing pattern.Abbreviations EOD electric organ discharge - HRP horseradish peroxidase - NMDA N-Methyl-D-Aspartate - PPN prepacemaker nucleus     

Electroreception,electrogenesis and electric signal evolution

Electroreception, the capacity to detect external underwater electric fields with specialised receptors, is a phylogenetically widespread sensory modality in fishes and amphibians. In passive electroreception, a capacity possessed by c. 16% of fish species, an animal uses low-frequency-tuned ampullary electroreceptors to detect microvolt-range bioelectric fields from prey, without the need to generate its own electric field. In active electroreception (electrolocation), which occurs only in the teleost lineages Mormyroidea and Gymnotiformes, an animal senses its surroundings by generating a weak (< 1 V) electric-organ discharge (EOD) and detecting distortions in the EOD-associated field using high-frequency-tuned tuberous electroreceptors. Tuberous electroreceptors also detect the EODs of neighbouring fishes, facilitating electrocommunication. Several other groups of elasmobranchs and teleosts generate weak (< 10 V) or strong (> 50 V) EODs that facilitate communication or predation, but not electrolocation. Approximately 1.5% of fish species possess electric organs. This review has two aims. First, to synthesise our knowledge of the functional biology and phylogenetic distribution of electroreception and electrogenesis in fishes, with a focus on freshwater taxa and with emphasis on the proximate (morphological, physiological and genetic) bases of EOD and electroreceptor diversity. Second, to describe the diversity, biogeography, ecology and electric signal diversity of the mormyroids and gymnotiforms and to explore the ultimate (evolutionary) bases of signal and receptor diversity in their convergent electrogenic–electrosensory systems. Four sets of potential drivers or moderators of signal diversity are discussed. First, selective forces of an abiotic (environmental) nature for optimal electrolocation and communication performance of the EOD. Second, selective forces of a biotic nature targeting the communication function of the EOD, including sexual selection, reproductive interference from syntopic heterospecifics and selection from eavesdropping predators. Third, non-adaptive drift and, finally, phylogenetic inertia, which may arise from stabilising selection for optimal signal-receptor matching.     

Primary afferent fibers conduct impulses in both directions under physiological stimulus conditions

Summary In electric fish of the family Mormyridae some primary afferent fibers conduct impulses not only from electroreceptors to the brain but also from the brain to the receptors. The efferent impulses may be elicited by electrical stimulation which is within the physiological range, i.e., by stimulation which is similar in amplitude and duration to the stimulation that is caused by the fish's own electric organ discharge. Afferent and efferent impulses in the same afferent fiber were identified by: simultaneously recording from a fiber at two different points, at the receptor and at the nerve trunk (Figs. 2C-H; 3B-D); by cutting the afferent fiber between the brain and the recording site as well as between the recording site and the periphery; and by intra-axonal recording from the afferent fiber near its entry into the brain (Fig. 4). The efferent impulses result from the central integration of a corollary discharge of the electric organ motor command with excitatory and inhibitory input from several different receptors near the one from which afferent impulses originate (Fig. 4). The centrally originating impulse may be capable of modifying the effect of signals originating in the periphery.Abbreviations ELLL electrosensory lateral line lobe - EOCD electric organ corollary discharge - EOD electric organ discharge - epsp excitatory postsynaptic potential - NPLL posterior lateral line nerve     

The mandibular common inhibitor system

Summary The mandibular common inhibitor neurones ofHomarus gammarus receive sensory input from a wide receptive field (Table 1, Figs. 2, 3) and from their symmetrical homologue (Ferrero and Wales, 1976).The CI system receives excitatory input from mandibular proprioceptors, with the notable exception of the mandibular muscle receptor organ, and its activity increases, during mandible opening and closing, towards the extremes of movement (Fig. 1). The output of CI neurones is usually coupled except during some high frequency bursts. Unilateral sensory input usually produces a coupled output. Electrical stimulation of a wide range of mandibular nerves (Table 2) has a similar effect and entrains the CI output at lower frequencies (Figs. 4, 5).Antidromic stimulation of a CI neurone causes excitation of its homologue but to a lower level of activity and without enhanced coupling. Even when the excitatory state is raised, by concurrent stimulation of a sensory nerve, the pathway activated by antidromic stimuli does not produce coupled activity at frequencies above 20 Hz (Fig. 8).Stimulation with single pulses will frequently produce short trains of impulses from the CI neurones (Figs. 6, 7) suggesting reciprocal excitation between the neurones.A model of the system based on current knowledge is presented.     

Behavioral responses of weakly electric fish to complex impedances

Summary Members of the family of African electric fish, Mormyridae, exhibit a novelty response, consisting of an acceleration in the rate of electric organ discharges (EODs), when faced with changes in feedback arising from their EODs. In this study, the novelty responses of three different species of mormyrids to shunts with different electrical characteristics were noted. The three species differed in the frequency contents of their EODs: two species had relatively high spectral frequencies in their EODs (>10 kHz), while the third species had only lower spectral frequencies (< 10 kHz). Primarily resistive shunts elicited novelty response accelerations in all three species, and the magnitudes of these responses, when normalized to the responses obtained for a shunt with no introduced resistance, were comparable for all three species. For primarily capacitive shunts, however, the magnitudes of the normalized responses were different for the three species: the two species with high spectral frequencies in their EODs showed larger normalized responses than the third species which had only low EOD spectral frequencies.The differences in species responses for capacitive shunts, and the similarities in species responses for resistive shunts, suggest that electric fish detect the complex impedance of objects in their near field environment: a circuit model consisting of a fish emitting discharges into the surrounding water, which can be shunted by a variable complex impedance, conforms well to the data. Thus, electrolocation is a frequency dependent sensory process, and this frequency dependency should be considered in any speculation about the adaptive value of different EOD waveforms.Abbreviation EOD electric organ discharge     

Communication in the weakly electric fish Sternopygus macrurus

There is a sexual dimorphism in the frequency of the quasi-sinusoidal electric organ discharge (EOD) of Sternopygus macrurus, with males, on average, an octave lower. EODs are detected by tuberous electroreceptor organs, which exhibit V-shaped frequency tuning with maximal sensitivity near the fish's own EOD frequency. This would seem to limit the ability of a fish to detect the EODs of opposite-sex conspecifics. However, electroreceptor tuning has always been based on single-frequency stimulation, while actual EOD detection involves the addition of a conspecific EOD to the fish's own. In the present study, recordings were made from single electroreceptive units while the fish were stimulated with pairs of sine waves: one (S1) representing the fish's own EOD added to a second (S2) representing a conspecific EOD. T unit response was easily predicted by assuming that the electroreceptor acts as a linear filter in series with a threshold-sensitive spike initiator. P unit response was more complex, and unexpectedly high sensitivity was found for frequencies of S2 well displaced from the fish's EOD frequency. For both P and T units, detection thresholds for S2 were much lower when added to S1, than when presented alone.     

Behavioral detection of electric signal waveform distortion in the weakly electric fish,Gnathonemus petersii

The novelty response of weakly electric mormyrids is a transient acceleration of the rate of electric organ discharges (EOD) elicited by a change in stimulus input. In this study, we used it as a tool to test whether Gnathonemus petersii can perceive minute waveform distortions of its EOD that are caused by capacitive objects, as would occur during electrolocation. Four predictions of a hypothesis concerning the mechanism of capacitance detection were tested and confirmed: (1) G. petersii exhibited a strong novelty response to computer-generated (synthetic) electric stimuli that mimic both the waveform and frequency shifts of the EOD caused by natural capacitive objects (Fig. 3). (2) Similar responses were elicited by synthetic stimuli in which only the waveform distortion due to phase shifting the EOD frequency components was present (Fig. 4). (3) Novelty responses could reliably be evoked by a constant amplitude phase shifted EOD that effects the entire body of the fish evenly, i.e., a phase difference across the body surface was lacking (Figs. 3, 4). (4) Local presentation of a phase-shifted EOD mimic that stimulated only a small number of electroreceptor organs at a single location was also effective in eliciting a behavioral response (Fig. 5).Our results indicate that waveform distortions due to phase shifts alone, i.e. independent of amplitude or frequency cues, are sufficient for the detection of capacitive, animate objects. Mormyrids perceive even minute waveform changes of their own EODs by centrally comparing the input of the two types of receptor cells within a single mormyromast electroreceptor organ. Thus, no comparison of differentially affected body regions is necessary. This shows that G. petersii indeed uses a unique mechanism for signal analysis, which is different from the one employed by gymnotiform wavefish.Abbreviations EOD electric organ discharge - p-p-amplitude peak-to-peak amplitude     

Shifts in frequency tuning of electroreceptors in androgen-treated mormyrid fish

Summary Several species of mormyrid electric fish have a sex difference in the pulse waveform of their electric organ discharge (EOD). Field studies in Gabon, West Africa have shown for one such species,Brienomyrus brachyistius (triphasic), that the sexually mature male EOD differs in shape and is nearly twice the duration of the EODs of females and juveniles. Fourier analysis reveals that differences in EOD duration correlate with those in the EOD power spectrum which has a peak at 0.3 kHz in males and 1.3 kHz in females and juveniles. We find a corresponding sex difference in the frequency tuning of at least one class of electroreceptors known as Knollenorgans. The average best or characteristic frequency of Knollenorgans is lower in males compared to females and juveniles. This correlates with a lower peak in the power spectrum of the male's pulse. When females are treated with gonadal androgens, their EODs increase 2–3 fold in duration, and the power spectra of their pulses are correspondingly lowered to match that of mature males. The average best frequency of Knollenorgans decreases by nearly 1 kHz which matches the downward shift of their EOD's power spectrum.For a second species ofBrienomyrus (sp. 2) which is commercially imported from Nigeria, we have not detected a sex difference in the power spectrum or duration of the EOD. The power spectrum peaks at about 4.2 kHz in males, females, and juveniles. Androgens, however, do cause a coincident downward shift in the average peak of the EOD power spectrum (from 4.2 to 1.3 kHz) and the average best frequency of Knollenorgans (from 2.3 to 1.4 kHz).Specimens ofBrienomyrus (sp. 2) that have been electrically silenced by surgical means are tuned, on the average, only 0.2 kHz higher than control animals. Silenced animals that have been treated with androgens are tuned, on the average, 0.2 kHz below controls. The results suggest that electroreceptor tuning is only partially modifiable during androgen treatment if the electroreceptors arenot being stimulated by an external electrical stimulus, i.e. the animal's own EOD. Since androgen treatment has a dramatic effect on receptor tuningonly in intact fish, it seems likely that retuning isnot due to a direct action of androgens on receptors, but rather due to the action of the principal electrical stimulus upon the receptors, i.e. the EOD. The implications of such results for the development of species and sex differences in electro-receptor tuning is discussed.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

The coding of signals in the electric communication of the gymnotiform fish Eigenmannia: From electroreceptors to neurons in the torus semicircularis of the midbrain

Summary In the context of aggression and courtship, Eigenmannia repeatedly interrupts its electric organ discharges (EODs) These interruptions (Fig. 1) contain low-frequency components as well as high-frequency transients and, therefore, stimulate ampullary and tuberous electroreceptors, respectively (Figs. 2, 3). Information provided by these two classes of receptors is relayed along separate pathways, via the electrosensory lateral line lobe (ELL) of the hindbrain, to the dorsal torus semicircularis (TSd) of the midbrain. Some neurons of the torus receive inputs from both types of receptors (Figs. 14, 15), and some respond predominantly to EOD interruptions while being rather insensitive to other forms of signal modulations (Figs. 12, 13). This high selectivity appears to result from convergence and gating of inputs from individually less selective neurons.Abbreviations CP central posterior thalamic nucleus - Df frequency difference between neighbor's EOD and fish's own - DPn dorsal posterior nucleus (thalamus) - EOD electric organ discharge - ELL electrosensory lateral line lobe - JAR jamming avoidance response - LMR lateral mesencephalic reticular formation - nE nucleus electrosensorius - nEb nucleus electrosensorius, beat-related area - nE nucleus electrosensorius, area causing rise of EOD frequency - nE nucleus electrosensorius, area causing fall of EOD frequency - nEar nucleus electrosensorius-acusticolateralis area - NPd nucleus praeeminentialis, pars dorsalis - PPn prepacemaker nucleus - PT pretectal nucleus - SE nucleus subelectrosensorius - TeO optic tectum - TSd dorsal (electrosensory) torus semicircularis - TSv ventral (mechano-sensory and auditory) torus semicircularis     

The course control system of beetles walking in an air-current field

The wind-orientated walk of carrion beetles Necrophorus humator F. was analysed under closed-loop conditions with a walking compensator and under openloop conditions with a paired tread wheel (Fig. 1).

Facilitation at crayfish neuromuscular junctions

Electrophysical recordings from opener muscle fibers in the crayfishProcambarus clarkii (Fig. 1) show that pre-synaptic facilitation at terminals of the single excitatory axon usually decays in a dual-exponential fashion after a single pulse or after a train of pulses (Figs. 2, 3, 7, 9), as has been reported for frog neuromuscular junctions (Mallart and Martin, 1967) and squid giant synapses (Charlton and Bittner, 1974, 1976). Furthermore, the second component of decay at crayfish synapses is associated with a break in the monotonic decay of the first component, a result which suggests that the decay of facilitation is not due to the simple diffusion of some substance (such as calcium) from specialized release sites.The growth of facilitation at all opener synapses during trains of equalinterval stimuli could not be predicted by assuming that each pulse contributed an equal amount of facilitation which summed linearly with that remaining from all previous stimuli (Figs. 4, 6; Table 2), as reported for synapses in frog and squid. During high frequency stimulation (>40 Hz), those terminals which facilitate dramatically (highF _e synapses) show much greater amounts of facilitation than that predicted by the linear summation model (Figs. 4, 8), whereas other terminals (lowF _e synapses) show much less facilitation than predicted (Fig. 6). The rate of growth of facilitation was often very constant at various stimulus rates in highF _e or mixed type synapses (Figs. 4, 8, 10)-a result not predicted by the linear summation model. Finally, when highF _e synapses were stimulated at different frequencies, the rate of growth of facilitation changed dramatically in a fashion not predictable using linear summation (Mallert and Martin, 1967) or power law (Linder, 1974) models.     

The echolocation and hunting behavior of the bat,Pipistrellus kuhli

Summary The echolocation and hunting behavior ofPipistrellus kuhli was studied in the field using multi-exposure photography synchronized with high-speed tape recordings. During the search phase, the bats used 8–12 ms signals with sweeps (sweep width 3–6 kHz) and pulse intervals near 100 ms or less often near 200 ms (Figs. 1 and 2). The bats seemed to have individual terminal frequencies that could lie between 35 and 40 kHz. The duty cycle of searching signals was about 8%. The flight speed of hunting bats was between 4.0 and 4.5 m/s. The bats reacted to insect prey at distances of about 70 to 120 cm. Given the flight speed, the detection distance was estimated to about 110 to 160 cm. Following detection the bat went into the approach phase where the FM sweep steepened (to about 60 kHz bandwidth) and the repetition rate increased (to about 30 Hz). The terminal phase or buzz, which indicates prey capture (or attempted capture), was composed of two sections. The first section contained signals similar to those in the approach phase except that the pulse duration decreased and the repetition rate increased. The second section was characterized by a sharp drop in the terminal frequency (to about 20 kHz) and by very short pulses (0.3 ms) at rates of up to 200 Hz (Figs. 1 and 3). Near the beginning of the buzz the bat prepared for capturing the prey by extending the wings and forming a tail pouch (Fig. 4). A pause of about 100 ms in sound emission after the buzz indicated a successful capture (Fig. 4). Pulse duration is discussed in relation to glint detection and detection distance. It is argued that the minimum detection distance can be estimated from the pulse duration as the distance where pulse-echo overlap is avoided.Abbreviations CF constant frequency - FM frequency modulated     

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).     

‘Communication’ in weakly electric fish, Gnathonemus petersii (Mormyridae) II. Interaction of electric organ discharge activities of two fish

The electric organ discharges (EODs) of pairs of weakly electric fish, Gnathonemus petersii, were simultaneously recorded to study the significance of the EODs as communication signals. In a 400-litre tank a larger fish (12 to 15 cm) was passively moved within a shelter tube toward a smaller specimen (6 to 9 cm), either in steps or a continuous move. The movement was stopped at that distance when at least one fish significantly lowered or ceased its EOD activity. From this ‘threshold interfish distance’ the spatial range of a ‘communication field’ was found to extend about 30 cm from the fish. At threshold distances an EOD frequency increase caused a temporary EOD activity cessation in the second fish. The spontaneous irregular EOD pattern of the fish displaying the increased EOD rate changed into a regular one with almost equal time intervals between fish pulses.