Inter‐annual variation in herring,Clupea harengus,and sprat,Sprattus sprattus,condition in the central Baltic Sea: what gives the tune?

The Baltic Sea ecosystem has undergone large changes during the last two decades, including a severe reduction in cod and herring biomass but, at the same time, a large increase in sprat abundance. The lower trophic levels of the Baltic Sea also changed due to environmental fluctuations, including variations in salinity and in volume of oxygenated water. In this apparently shifting environment, the conditions of herring and sprat have undergone large inter-annual variations during the past 15–20 years. In this study, we explore how abiotic factors (i.e. salinity and temperature) and biotic factors (biomass of the copepods Pseudocalanus elongatus , Temora longicornis , Acartia spp. and of cladocerans as well as clupeid abundance) in different seasons (May and August) affect clupeid body condition. Our analyses suggest that data of zooplankton biomass and abiotic factors in August have higher predictive power than May data. Although our analysis suggests that salinity (a bottom-up process) has an effect on sprat condition, total abundance of clupeids (a top-down process) is by far the most significant predictor of both herring and sprat condition. The strong correlation between clupeid abundance and total zooplankton biomass points to food competition and to top-down control by herring and sprat on common food resources. Furthermore, clupeid condition co-varied with the changes in the weight of zooplankton in the stomachs, which further suggest food competition being the main mechanism behind the changes in clupeid condition during the last two decades. Hence, our results are not in agreement with most of the current literature that has suggested that clupeid growth is regulated by environmentally mediated bottom-up processes acting on the abundance of copepods. This is, to our knowledge, the first evidence of food resources mediated density-dependent fish growth in a large marine ecosystem.     

Changes in haematological parameters in relation to prey switching in a wild population of harbour seals

1. Previous studies have found marked inter-annual variation in winter food availability, diet composition and body condition in a population of harbour seals ( Phoca vitulina L.) in Northeast Scotland. This study aimed to determine whether there were other physiological consequences of prey switching by comparing haematological parameters in years when the clupeids herring and sprat dominated the diet and in years when seals switched to alternative prey.
2. There were significant differences in leukocyte and erythrocyte parameters in relation to diet composition. In contrast, indices of body condition did not explain the variation in haematological parameters, suggesting that the observed changes did not result from differences in the energetic content of the prey.
3. Leukocyte counts were significantly higher after 'good' clupeid years, although the differences in mean counts were small. Such differences could have resulted either from immuno-suppression, for example because of differences in prey nutrient or contaminant levels, or from differences in the pathogen challenge resulting from geographical variations in water quality.
4. The differences in erythrocyte parameters were more marked, and there was evidence of widespread macrocytic anaemia when seals switched from clupeids to alternative prey. Such differences could result either from acclimation, as a result of prey-specific foraging strategies, or from differences in the nutritional quality of prey.
5. These results indicate that generalist predators such as the harbour seal may exhibit physiological responses to changes in the composition of their diet. These data highlight the need to consider the long-term physiological effects of variations in food availability on the population dynamics of generalist marine top predators. In particular, it is hypothesized that fish-induced anaemia may be responsible for declines in certain pinniped populations.     

Hearing sensitivity in two black bass species using the auditory brainstem response approach

Recently, several bioacoustic studies have focused on the red eye bass (Micropterus coosae). One of these studies documented sound production, while the other played back sounds produced by prey items in order to determine their attractiveness to M. coosae. Surprisingly, the hearing ability of fishes in the genus Micropterus has received very little attention. The need for audiograms describing hearing in Micropterus is apparent. This study utilized the auditory brainstem response (ABR) approach to determine hearing sensitivity in terms of both sound pressure level (SPL) and particle acceleration in two black bass species, the red eye bass (M. coosae) and the Alabama bass (M. henshalli). Audiograms produced in this study expressed in both SPL and particle acceleration showed a positive relationship between hearing threshold and frequency. Micropterus coosae was most sensitive to frequencies that overlap with the peak frequencies of their vocalizations, and the vocalizations of a prey species, Cyprinella trichroistia. Bass hearing sensitivities at lower frequencies, measured in terms of particle acceleration, were similar to several sciaenid species.     

Abundance,structure, growth and origin of inshore clupeid populations of the west coast of Scotland

The abundance, structure, growth, and origin of clupeid populations in inshore waters of the west coast of Scotland were studied from April 1970 to October 1972. Clupeid populations in the area comprise mainly young fish. 0-group autumn-spawned herring, probably of Minch origin, move into the area about April and spring-spawned ones (Clyde origin) about June. The timing and the body length at which each group arrives in the area during the different years is the same. After the initial immigration, the distribution of both 0-group clupeids becomes localized.Herring populations in the sea-lochs and associated inshore waters are mainly 0-group fish, which are replaced each year by a new incoming brood. The sprat populations of the sea-lochs are dominated by the 0-group; in the more ‘open’ areas the populations comprise older individuals. Year-class distribution in the ‘open’ areas resemble that of the commercial fishery.The rate of increase of 0-group autumn-spawned herring is 3.68 mm/week and that of spring-spawned 0-group herring is 2.83 mm/week. The curves of growth of 0-group sprats of the 1970 and 1971 year-classes are different; the rate of increase in length, however, averages 3.55 mm/week for both year-classes and the differences are not significant.In sprats, after their first year of life a rapid increase in length takes place in the spring. This increase is thought to enable the majority of the population to reach the minimum size (88–90 mm) for initial gonadal maturation and thereby make them capable of reproducing in their second year of life.     

Keeping up with bats: dynamic auditory tuning in a moth

Many night-flying insects evolved ultrasound sensitive ears in response to acoustic predation by echolocating bats . Noctuid moths are most sensitive to frequencies at 20-40 kHz , the lower range of bat ultrasound . This may disadvantage the moth because noctuid-hunting bats in particular echolocate at higher frequencies shortly before prey capture and thus improve their echolocation and reduce their acoustic conspicuousness . Yet, moth hearing is not simple; the ear's nonlinear dynamic response shifts its mechanical sensitivity up to high frequencies. Dependent on incident sound intensity, the moth's ear mechanically tunes up and anticipates the high frequencies used by hunting bats. Surprisingly, this tuning is hysteretic, keeping the ear tuned up for the bat's possible return. A mathematical model is constructed for predicting a linear relationship between the ear's mechanical stiffness and sound intensity. This nonlinear mechanical response is a parametric amplitude dependence that may constitute a feature common to other sensory systems. Adding another twist to the coevolutionary arms race between moths and bats, these results reveal unexpected sophistication in one of the simplest ears known and a novel perspective for interpreting bat echolocation calls.     

Hearing in the African lungfish (Protopterus annectens): pre-adaptation to pressure hearing in tetrapods?

Lungfishes are the closest living relatives of the tetrapods, and the ear of recent lungfishes resembles the tetrapod ear more than the ear of ray-finned fishes and is therefore of interest for understanding the evolution of hearing in the early tetrapods. The water-to-land transition resulted in major changes in the tetrapod ear associated with the detection of air-borne sound pressure, as evidenced by the late and independent origins of tympanic ears in all of the major tetrapod groups. To investigate lungfish pressure and vibration detection, we measured the sensitivity and frequency responses of five West African lungfish (Protopterus annectens) using brainstem potentials evoked by calibrated sound and vibration stimuli in air and water. We find that the lungfish ear has good low-frequency vibration sensitivity, like recent amphibians, but poor sensitivity to air-borne sound. The skull shows measurable vibrations above 100 Hz when stimulated by air-borne sound, but the ear is apparently insensitive at these frequencies, suggesting that the lungfish ear is neither adapted nor pre-adapted for aerial hearing. Thus, if the lungfish ear is a model of the ear of early tetrapods, their auditory sensitivity was limited to very low frequencies on land, mostly mediated by substrate-borne vibrations.     

褐菖鲉的听觉阈值研究

利用听觉诱发电位记录技术研究了褐菖鲉(Sebasticus marmoratus)的听觉阈值。通过采用听觉生理系统记录和分析了8尾褐菖鲉对频率范围在100—1000 Hz的7种不同频率的声音刺激的诱发电位反应。结果表明, 褐菖鲉的听觉阈值在整体上随着频率增加而增加, 对100—300 Hz的低频声音信号敏感, 最敏感频率为150 Hz, 对应的听觉阈值为70 dB re 1 μPa。褐菖鲉的听觉敏感区间与其发声频率具有较高的匹配性, 表明其声讯交流的重要性。同时, 人为低频噪声可能对其声讯交流造成影响。     

3-D-orientation with the octavolateralis system.

Fish detect and localize a sound source with inner ear receptors and with the mechanosensory lateral line. The inner ear of fish is sensitive to the water displacements caused by sound waves through a direct, inertial response by hair cell epithelia of the ear. Hearing specialists, such as goldfish and herring, have accessory peripheral structures that provide additional sensitivity to the pressure component of a sound wave. While the inner ear of fish responds to the whole body motions caused by sound waves and--in case of hearing specialists--to sound pressure, the lateral line is only sensitive to water motions relative to the surface of the fish and to local pressure gradients. Using lateral line and/or acoustic input, some fish can determine the direction and the distance to a sound source. Most likely they do so by exploiting some of the mechanisms described in this paper. Piscivorous fish may use lateral line input to detect the wakes caused by swimming fish. Even in the absence of light catfish, for instance, can follow a 10 s old, three-dimensional wake left by a prey fish over distances up to 55 prey-body length.     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

The ontogenetic development of auditory sensitivity, vocalization and acoustic communication in the labyrinth fish Trichopsis vittata

The anabantoid fish Trichopsis vittata starts vocalizing as 8-week-old juveniles. In order to determine whether juveniles are able to detect conspecific sounds, hearing sensitivities were measured in six size groups utilizing the auditory brainstem response-recording technique. Results were compared to sound pressure levels and spectra of sounds recorded during fighting. Auditory evoked potentials were present in all size groups and complete audiograms were obtained starting with 0.18 to 0.30 g juveniles. Auditory sensitivity during development primarily increased between 0.8 kHz and 3.0 kHz. The most sensitive frequency within this range shifted from 2.5 kHz to 1.5 kHz, whereas thresholds decreased by 14 dB. Sound production, on the other hand, started at 0.1 g and sound power spectra at dominant frequencies increased by 43 dB, while dominant frequencies shifted from 3 kHz to 1.5 kHz. Comparisons between audiograms and sound power spectra in similar-sized juveniles revealed no clear match between most sensitive frequencies and dominant frequencies of sounds. This also revealed that juveniles cannot detect conspecific sounds below the 0.31 to 0.65 g size class. These results indicate that auditory sensitivity develops prior to the ability to vocalize and that vocalization occurs prior to the ability to communicate acoustically.     

Better than fish on land? Hearing across metamorphosis in salamanders

Early tetrapods faced an auditory challenge from the impedance mismatch between air and tissue in the transition from aquatic to terrestrial lifestyles during the Early Carboniferous (350 Ma). Consequently, tetrapods may have been deaf to airborne sounds for up to 100 Myr until tympanic middle ears evolved during the Triassic. The middle ear morphology of recent urodeles is similar to that of early ‘lepospondyl’ microsaur tetrapods, and experimental studies on their hearing capabilities are therefore useful to understand the evolutionary and functional drivers behind the shift from aquatic to aerial hearing in early tetrapods. Here, we combine imaging techniques with neurophysiological measurements to resolve how the change from aquatic larvae to terrestrial adult affects the ear morphology and sensory capabilities of salamanders. We show that air-induced pressure detection enhances underwater hearing sensitivity of salamanders at frequencies above 120 Hz, and that both terrestrial adults and fully aquatic juvenile salamanders can detect airborne sound. Collectively, these findings suggest that early atympanic tetrapods may have been pre-equipped to aerial hearing and are able to hear airborne sound better than fish on land. When selected for, this rudimentary hearing could have led to the evolution of tympanic middle ears.     

Auditory sensitivity in settlement-stage larvae of coral reef fishes

The larval phase of most species of coral reef fishes is spent away from the reef in the pelagic environment. At the time of settlement, these larvae need to locate a reef, and recent research indicates that sound emanating from reefs may act as a cue to guide them. Here, the auditory abilities of settlement-stage larvae of four species of coral reef fishes (families Pomacentridae, Lutjanidae and Serranidae) and similar-sized individuals of two pelagic species (Carangidae) were tested using an electrophysiological technique, auditory brainstem response (ABR). Five of the six species heard frequencies in the 100–2,000 Hz range, whilst one carangid species did not detect frequencies higher than 800 Hz. The audiograms of the six species were of similar shape, with best hearing at lower frequencies between 100 and 300 Hz. Strong within-species differences were found in hearing sensitivity both among the coral reef species and among the pelagic species. Larvae of the coral reef species had significantly more sensitive hearing than the larvae of the pelagic species. The results suggest that settlement-stage larval reef fishes may be able to detect reef sounds at distances of a few 100 m. If true hearing thresholds are lower than ABR estimates, as indicated in some comparisons of ABR and behavioural methods, the detection distances would be much larger.     

Detection of low‐frequency tones and whale predator sounds by the American sand lance Ammodytes americanus

Auditory evoked potentials (AEP) were used to measure the hearing range and auditory sensitivity of the American sand lance Ammodytes americanus. Responses to amplitude‐modulated tone pips indicated that the hearing range extended from 50 to 400 Hz. Sound pressure thresholds were lowest between 200 and 400 Hz. Particle acceleration thresholds showed an improved sensitivity notch at 200 Hz but not substantial differences between frequencies and only a slight improvement in hearing abilities at lower frequencies. The hearing range was similar to Pacific sand lance Ammodytes personatus and variations between species may be due to differences in threshold evaluation methods. AEPs were also recorded in response to pulsed sounds simulating humpback whale Megaptera novaeangliae foraging vocalizations termed megapclicks. Responses were generated with pulses containing significant energy below 400 Hz. No responses were recorded using pulses with peak energy above 400 Hz. These results show that A. americanus can detect the particle motion component of low‐frequency tones and pulse sounds, including those similar to the low‐frequency components of megapclicks. Ammodytes americanus hearing may be used to detect environmental cues and the pulsed signals of mysticete predators.     

Comparative otolith morphology of clupeids from the Iranian brackish and marine resources (Teleostei: Clupeiformes)

The diagnostic features of otolith morphology were provided for the clupeid fishes of the Iranian brackish and marine resources to be used as diagnostic features for the identification of clupeids diversity in these regions. Fish individuals belong to 20 species belong to 13 genera, and four families were collected from the Caspian Sea, the Persian Gulf and the Makran zone of the Oman Sea. Overall, seven otolith morphotypes were distinguished, that is lanceolated (45%), fusiform (20%), boot-like (15%) and clamp-like, pentagonal, elliptic and rectangular (each 5%, respectively). The univariate analysis showed that all variables except shape index [sulcus area (SS)/otolith area (OS)] and ROx (roundness) were significantly different among the clupeid species. The interspecific relationships of the otoliths were analysed based on the data of otolith morphology and otolith shape indices. Both dendrograms showed somehow an obvious separation among the studied species. However, the interspecific relationships in the dendrogram based on the otolith morphology have a better resolution. These phenotypic relationships based on otolith morphology among the studied clupeids are largely consistent with the previous hypothesis on the systematics of these fishes and emphasized that the morphological and morphometric features of the otolith, which are highlighted in this study, could be adequately used as diagnostic features for the identification of clupeids diversity.     

Listening for males and bats: spectral processing in the hearing organ of <Emphasis Type="Italic">Neoconocephalus bivocatus</Emphasis> (Orthoptera: Tettigoniidae)

Tettigoniids use hearing for mate finding and the avoidance of predators (mainly bats). Using intracellular recordings, we studied the response properties of auditory receptor cells of Neoconocephalus bivocatus to different sound frequencies, with a special focus on the frequency ranges representative of male calls and bat cries. We found several response properties that may represent adaptations for hearing in both contexts. Receptor cells with characteristic frequencies close to the dominant frequency of the communication signal were more broadly tuned, thus extending their range of high sensitivity. This increases the number of cells responding to the dominant frequency of the male call at low signal amplitudes, which should improve long distance call localization. Many cells tuned to audio frequencies had intermediate thresholds for ultrasound. As a consequence, a large number of receptors should be recruited at intermediate amplitudes of bat cries. This collective response of many receptors may function to emphasize predator information in the sensory system, and correlates with the amplitude range at which ultrasound elicits evasive behavior in tettigoniids. We compare our results with spectral processing in crickets, and discuss that both groups evolved different adaptations for the perceptual tasks of mate and predator detection.     

The auditory system of blood-sucking mosquito females (Diptera, Culicidae): Acoustic perception during flight simulation

Mosquitoes hear with their plumose antennae which respond to the air movement caused by sound propagation and conduct vibrations to the Johnston’s organ located at the base of each antenna. Each of the two Johnston’s organs contains several tens of thousands mechanosensory cells which detect the displacements of the flagellum and transform them into electric potentials. Hearing plays a very important role in the reproductive behavior of the male mosquitoes. At the same time, our knowledge of hearing in female mosquitoes is very limited and its functional significance is obscure. In this study we measured the auditory sensitivity of female mosquitoes and investigated how the flight conditions affect their hearing. We studied mosquitoes of three species: Anopheles messeae, Aedes excrucians, and Culex pipiens pipiens. The neuronal responses were recorded with a glass microelectrode from the antennal nerve and the deutocerebral interneurons. Stimulation was applied in two modes: (1) the main stimulus against the background of flight simulation (strong vibration with the typical wingbeat frequency of a given mosquito species) and (2) only the main stimulus without the background stimulation. During the flight simulation, females demonstrated an increased sensitivity to frequencies below 200 Hz. The mean auditory receptor threshold at 80–120 Hz was 45 dB, which was 8 dB lower than that without flight simulation. An additional zone of increased sensitivity was also found at frequencies higher than the simulated wingbeat frequency (the so-called image channel). Our analysis of frequency tuning curves measured from the receptors and auditory interneurons shows that mosquito auditory neuronal complex consists of several subsystems which have different frequency tuning parameters, and suggests the possibility of spectral analysis of sounds. Three hypotheses could be proposed on the function of hearing in female mosquitoes: (1) predator avoidance, (2) detection of moving prey, and (3) intraspecific communication. Each of the hypotheses involves the ability to analyze the sound frequency spectrum and subsequent signal recognition.     

Aerial hearing thresholds and detection of hearing loss in male California sea lions (Zalophus californianus) using auditory evoked potentials

Auditory evoked potential (AEP) measurements are useful for describing the variability of hearing among individuals in marine mammal populations, an important consideration in terms of basic biology and the design of noise mitigation criteria. In this study, hearing thresholds were measured for 16 male California sea lions at frequencies ranging from 0.5 to 32 kHz using the auditory steady state‐response (ASSR), a frequency‐specific AEP. Audiograms for most sea lions were grossly similar to previously reported psychophysical data in that hearing sensitivity increased with increasing frequency up to a steep reduction in sensitivity between 16 and 32 kHz. Average thresholds were not different from AEP thresholds previously reported for male and female California sea lions. Two sea lions from the current study exhibited abnormal audiograms: a 26‐yr‐old sea lion had impaired hearing with a high‐frequency hearing limit (HFHL) between 8 and 16 kHz, and an 8‐yr‐old sea lion displayed elevated thresholds across most tested frequencies. The auditory brainstem responses (ABRs) for these two individuals and an additional 26‐yr‐old sea lion were aberrant compared to those of other sea lions. Hearing loss may have fitness implications for sea lions that rely on sound during foraging and reproductive activities.     

Ultrasound sensitive neurons in the cricket brain

1. The aim of this study was to identify neurons in the brain of the cricket, Teleogryllus oceanicus, that are tuned to high frequencies and to determine if these neurons are involved in the pathway controlling negative phonotaxis. In this paper we describe, both morphologically and physiologically, 20 neurons in the cricket brain which are preferentially tuned to high frequencies. 2. These neurons can be divided into two morphological classes: descending brain interneurons (DBINs) which have a posteriorly projecting axon in the circumesophageal connective and local brain neurons (LBNs) whose processes reside entirely within the brain. All the DBINs and LBNs have processes which project into one common area of the brain, the ventral brain region at the border of the protocerebrum and deutocerebrum. Some of the terminal arborizations of Int-1, an ascending ultrasound sensitive interneuron which initiates negative phonotaxis, also extend into this region. 3. Physiologically, ultrasonic sound pulses produce 3 types of responses in the DBINs and LBNs. (1) Seven DBINs and 6 LBNs are excited by ultrasound. (2) Ongoing activity in one DBIN and 5 LBNs is inhibited by ultrasound, and (3) one cell, (LBN-ei), is either excited or inhibited by ultrasound depending on the direction of the stimulus. 4. Many of the response properties of both the DBINs and LBNs to auditory stimuli are similar to those of Int-1. Specifically, the strength of the response, either excitation or inhibition, to 20 kHz sound pulses increases with increasing stimulus intensity, while the response latency generally decreases. Moreover, the thresholds to high frequencies are much lower than to low frequencies. These observations suggest that the DBINs and LBNs receive a majority of their auditory input from Int-1. However, the response latencies and directional sensitivity of only LBN-ei suggest that it is directly connected to Int-1. 5. The response of only one identified brain neuron, DBIN8, which is inhibited by 20 kHz sound pulses, is facilitated during flight compared to its response at rest. This suggests that suppression of activity in DBIN8 may be associated with ultrasound-induced negative phonotactic steering responses in flying crickets. The other DBINs and LBNs identified in this paper may also play a role in negative phonotaxis, and possibly in other cricket auditory behaviors influenced by ultrasonic frequencies.     

Ultrasound hearing and male–male communication in Australian katydids (Tettigoniidae: Zaprochilinae) with sexually dimorphic ears

The genus Kawanaphila (Tettigoniidae: Zaprochilinae) is unusual among the Tettigoniidae in the possession of sexually dimorphic auditory organs. We examined the auditory system and acoustic behaviour of two previously unstudied species in this genus to test whether reduced hearing in males is consistently associated with reduced male–male competition. Kawanaphila yarraga (Rentz, 1993) and K. mirla (Rentz, 1993) are both sexually dimorphic with respect to their auditory system, but to different degrees. Males of both species produce songs consisting of trains of brief (< 1 ms) pure-tone sound pulses at ultrasonic frequencies (K. yarraga, 40 kHz;K. mirla, 70 kHz). In both species, female hearing is more sensitive than that of males by 10 dB. In addition, male K. mirla are most sensitive at lower frequencies than females. Male and female K. yarraga differed only in sensitivity, not in tuning. The two species also differ in their degree of sexual dimorphism in auditory anatomy. Kawanaphila mirla males lack some auditory specializations of the prothoracic tracheal system, which are present in the normal tettigoniid condition in females. In K. yarraga males these structures are present, but reduced in size relative to females. The acoustic behaviour of males of the two species is consistent with this pattern of relative auditory sensitivity. Males of both species interact acoustically by altering the timing of their sound output to synchronize with neighbouring males. However, K. mirla males only interact in this way over very short distances (< 5 m), whereas K. yarraga males interact with neighbours up to at least 10 m distant. These results indicate that, although males of the two species differ in hearing sensitivity, the nature of their responses to conspecific calls are similar to one another and to those of other acoustic insects. This suggests that acoustically mediated male–male competition may be maintained even while selection favours a reduction in male auditory sensitivity.     

Escape manoeuvres of schooling Clupea harengus

The escape behaviour of schooling herring startled by an artificial sound stimulus was observed by means of high speed video filming. Response latencies showed two distinct peaks, at 30 ms and c . 100 ms. Escape responses belonging to the two latency groups showed different turning rates during the first stage of the response, and showed different escape trajectories. We suggest that long latency escapes may be responses to startled neighbours or simply weak responses to the sound stimulus. In addition, the different contraction rates during the C-bend formation seen in the two latency groups may imply differences in the neuronal commands. The escape responses of herring were directed away from the stimulus more often than towards it (88% of the total). These away responses were more common in long latency responses, suggesting that the latter enable herring to be more accurate in discerning the direction of the threat. Startled fish contracting their body towards the stimulus (performing a towards response) appear to correct their escape course, since their escape trajectory distribution is non-uniformty distributed around 360° and directed away from the stimulus. We hypothesize that when herring are schooling, the ability of each fish to correct its trajectory following turns towards the stimulus is enhanced.