Is there a fish alarm cue? Affirming evidence from a wild study

Chemical alarm cues released from injured tissue are not released under any other context and therefore reliably inform nearby prey of the presence of a predator. Laboratory and field studies have demonstrated that most aquatic taxa show antipredator responses to chemical alarm cues. Ostariophysan fish (e.g. minnows) possess specialized skin cells that contain an alarm chemical. Magurran et al. (1996, Proceedings of the Royal Society of London, Series B,263, 1551-1556) were the first to use underwater video to carefully document the behavioural response of free-ranging wild populations of minnows to minnow alarm cues. They found no evidence of an antipredator response, and challenged the assumption that the contents of these cells indicate risk in the field. They proposed that alarm responses are context dependent in that they are an artefact of enclosed environments such as laboratory aquaria and field traps. Here, we repeat their experiment on free-swimming field populations of littoral fish and report a significant decrease in the number of fish in areas where chemical alarm cues of blacknose shiners, Notropis heterolepis (Ostariophysi: Cyprinidae) were released. The effect of these chemical cues was equal in magnitude to the effect of the presentation of a model predator. The response to the approach of a model predator (visual cue) was intensified by pre-exposure to chemical alarm cues. We corroborated this interaction between chemical and visual indicators of predation risk in a laboratory study using glowlight tetras, Hemigrammus erythrozonus (Ostariophysi: Characidae). Response to the visual stimulus of a predator was significantly intensified by previous exposure to conspecific chemical alarm cues. We conclude that ostariophysan skin indeed contains an alarm cue that (1) informs nearby prey of imminent predation risk, (2) induces some form of antipredator behaviour in most contexts, and (3) affects subsequent behavioural responses to stimuli in other sensory modalities.     

Motion, not Shape, Facilitates Association of Predation Risk with Novel Objects by Fathead Minnows (Pimephales promelas)

Injury‐released chemical cues are reliable indicators of predation risk among many aquatic taxa. When a novel, neutral stimulus is presented in tandem with chemical cues from an injured conspecific, an association is formed between the novel stimulus and apparent risk. Learned recognition of predation risk is well documented for fathead minnows, Pimephales promelas. When minnows detect alarm cues in nature they are also potentially exposed to multiple environmental stimuli, few of which are likely to be relevant indicators of risk. How do minnows discern among candidate stimuli potentially associated with predation risk? Two possibilities are shape and motion. In this study, individual piscivore‐naïve minnows were presented simultaneously with conspecific chemical alarm cues and two stimulus objects. One object was a darkened tube with its long axis in the horizontal plane (fish‐like). The second object was a black disk. Following introduction of chemical alarm cues, one of the objects was raised and lowered repeatedly. After a single conditioning trial, minnows associated risk significantly more with the previously moving object than the previously stationary object, as indicated by reduced activity. Object shape had no significant effect on response intensity in test trials. Our data suggest that minnows have been selected to form aversive responses to moving objects at a site of recent predation because movement is a more predictable indicator of predator identity than shape.     

Assessment of local predation risk: the role of subthreshold concentrations of chemical alarm cues

The ability to accurately assess local predation risk is criticalto prey individuals, as it allows them to maximize threat-sensitivetrade-offs between predator avoidance and other fitness relatedactivities. A wide range of taxonomically diverse prey (includingmany freshwater fishes) relies on chemical alarm cues (alarmpheromones) as their primary information source for local riskassessment. However, the value of chemical alarm cues has beenquestioned due to the availability of additional sensory inputs(i.e., visual cues) and the lack of an overt antipredator responseunder conditions of low perceived risk. In this paper, we testthe hypothesis that chemical alarm cues at concentrations belowthe point at which they elicit an overt behavioral responsefunction to increase vigilance towards other sensory modalities(i.e., visual alarm cues). Shoals of glowlight tetras (Hemigrammuserythrozonus) exposed to the subthreshold concentration of hypoxanthine-3-N-oxide(the putative Ostariophysan alarm pheromone) did not exhibitan overt antipredator response in the absence of secondary visualcues (not different than the distilled water control). However,when exposed to the sight of a visually alarmed conspecific,they significantly increased the intensity of their antipredatorresponse (not different from shoals exposed to the suprathresholdalarm cue). This study demonstrates that prey may benefit fromresponding to low concentration alarm cues by increasing vigilancetowards secondary cues during local risk assessment, even inthe absence of an overt behavioral response. By increasing vigilancetowards secondary risk assessment cues in the presence of alow risk chemical cue, individuals are likely able to maximizethe threat-sensitive trade-offs between predator avoidance andother fitness related activities.     

Chemical Predator Inspection in a Characin Fish (Hemigrammus erythrozonus, Characidae, Ostariophysi): The Effects of Mixed Predator Diets

Many prey organisms will approach (inspect) potential predators, primarily to assess local risk of predation. It has been demonstrated that Ostariphysan prey fishes can detect conspecific alarm pheromones in the diet of potential predators and use this chemical information to reduce their risk of predation while still gaining significant benefits associated with predator inspection. We conducted the current study to examine the possible effects of mixed diets on the use of these chemical predator diet cues during inspection visits. Shoals of four glowlight tetras ( Hemigrammus erythrozonus ) were exposed to Jack Dempsey cichlids ( Cichlasoma octofaciatum ) which had been fed diets consisting of: 100% tetras (with alarm pheromone); 75% tetra, 25% swordtail ( Xiphophorus helleri , which lack a recognizable alarm pheromone); 25% tetra, 75% swordtail; or 100% swordtails. Tetras significantly increased their anti-predator behaviour in response to predators fed 100% tetra or the two mixed predator diets, but not when exposed to predators fed a 100% swordtail diet. Likewise, we observed significant differences in inspection behaviour. Tetras took longer to initiate an inspection, inspected in smaller groups and directed a greater proportion of inspection visits towards the tail region of the predator when it had been fed 100% tetra or either of the two mixed prey diets. We found no significant differences in either anti-predator or inspection behaviour among the three diet treatments containing tetras. These data strongly suggest that glowlight tetras are capable of detecting relatively small amounts of conspecific alarm pheromone in the diet of potential predators and that they modify their behaviour based on the presence or absence of these cues.     

Who dares, learns: chemical inspection behaviour and acquired predator recognition in a characin fish

Individuals that dare approach predators (predator inspection behaviour) may benefit by acquiring information regarding the potential threat of predation. Although information acquisition based on visual cues has been demonstrated for fish, it is unknown whether fish will inspect predators on the basis of chemical cues or whether such inspection behaviour results in information acquisition. Here, we first ascertained whether predator inspection behaviour can be mediated by chemical cues from predators by exposing groups of predator-naive glowlight tetras (Hemigrammus erythrozonus) to the chemical cues of a potential fish predator (convict cichlid Cichlasoma nigrofasciatum) that had been fed either tetras (which possess an alarm pheromone) or swordtails (Xiphophorus helleri, which lack Ostariophysan alarm pheromones). Tetras showed a significant increase in antipredator behaviour when exposed to the tetra-diet cue, but not when exposed to the swordtail-diet cue. Chemically mediated predator inspection behaviour was also affected. Both the latency to inspect and the minimum approach distance to the predator significantly increased, and the mean number of inspectors per predator inspection visit significantly decreased when tetras were exposed to the tetra-diet versus the swordtail-diet chemical cues. We then examined a potential benefit associated with chemically mediated predator inspection behaviour. Only tetras that were initially exposed to the tetra-diet cue and that had inspected the predator acquired the visual recognition of a convict cichlid as a predation threat. Our results thus demonstrate that (1) predator inspection behaviour in the glowlight tetra can be initiated by chemical cues, (2) chemically mediated inspection behaviour is affected by the presence of alarm pheromone, and (3) inspectors benefit by acquiring the recognition of novel predators. Copyright 1999 The Association for the Study of Animal Behaviour.     

Predator Inspection Behaviour in a Characin Fish: an Interaction between Chemical and Visual Information?

Recent evidence suggests that predator inspection behaviour by Ostariophysan prey fishes is regulated by both the chemical and visual cues of potential predators. In laboratory trials, we assessed the relative importance of chemical and visual information during inspection visits by varying both ambient light (visual cues) and predator odour (chemical cues) in a 2 × 2 experimental design. Shoals of glowlight tetras (Hemigrammus erythrozonus) were exposed to a live convict cichlid (Archocentrus nigrofasciatus) predator under low (3 lux) or high (50 lux) light levels and in the presence of the odour of a cichild fed tetras (with an alarm cue) or swordtails (Xiphophorus helleri, with an alarm cue not recognized by tetras). Tetras exhibited threat‐sensitive inspection behaviour (increased latency to inspect, reduced frequency of inspection, smaller inspecting group sizes and increased minimum approach distance) towards a predator paired with a tetra‐fed diet cue, regardless of light levels. Similar threat‐sensitive inspection patterns were observed towards cichlids paired with a swordtail‐fed diet cue only under high light conditions. Our data suggest that chemical cues in the form of prey alarm cues in the diet of the predator, are the primary source of information regarding local predation risk during inspection behaviour, and that visual cues are used when chemical information is unavailable or ambiguous.     

The Effects of Density on the Learned Recognition of Heterospecific Alarm Cues

Numerous species, both aquatic and terrestrial, use alarm cues to mediate predation risk. These cues may be either intentionally or inadvertently released, and may be received by either conspecifics or heterospecifics. In aquatic systems, alarm cues are often chemical in nature and are released when an organism is disturbed or damaged by a predator. In some cases the recognition of alarm cues from conspecifics, or closely related heterospecifics, is innate, while the recognition of alarm cues from distantly related species must be learned. Many studies have documented the use of heterospecific alarm cues, but few have explored the manner in which these cues come to be recognized as an indication of predation. In the current study, we examined the fathead minnow (Pimephales promelas)/brook stickleback (Culaea inconstans) alarm system. We tested the effect of density on the ability of minnows to learn to recognize stickleback alarm cues as a threat. We hypothesized that the ability of minnows to learn to recognize stickleback alarm cues should increase with increasing stickleback density because there would be more opportunity for minnows to associate the heterospecific alarm cue with the threat. To test this hypothesis we stocked minnows into large outdoor pools with no stickleback, low numbers of stickleback, or high numbers of stickleback. All pools contained a predator (pike, Esox lucius) known to the minnows. Following a 14 d conditioning period, minnows were tested for a response to skin extract from stickleback, minnow, and an unknown heterospecific (swordtail, Xiphophorus helleri). Minnows from pools with large numbers of stickleback learned to respond to stickleback alarm cues while minnows from pools with low numbers of stickleback, or no stickleback, did not.     

Acquired Recognition of Chemical Stimuli from Pike,Esox lucius,by Brook Sticklebacks,Culaea inconstans (Osteichthyes,Gasterosteidae)

In this study we test whether brook sticklebacks (Culaea inconstans) can acquire predator recognition through releaser-induced recognition learning, i.e. simultaneous exposure to aversive ('releasing') stimuli and neutral stimuli causing learned aversion to the neutral stimuli. We exposed wild-caught pike-naive brook sticklebacks (collected from a creek containing fathead minnows, Pimephales promelas, but not pike, Esox lucius) to chemical stimuli from pike that were mixed with brook stickleback skin extract, fathead minnow skin extract, or a control of distilled water. In subsequent tests 2 d later, when only pike stimuli were presented, sticklebacks conditioned with stickleback skin extract and fathead minnow skin extract exhibited antipredator behaviour (i.e. increased schooling and movement toward the substrate), while those conditioned with distilled water did not. Sticklebacks conditioned with stickleback skin extract responded to pike with a more intense response, in terms of movement toward the substrate, than those conditioned with fathead minnow skin extract, suggesting that conspecific skin extract may be a stronger stimulus than heterospecific skin extract for learning recognition of predators. To our knowledge this is the first study to demonstrate that an acanthopterygian fish can acquire predator recognition through the pairing of conspecific alarm pheromone with the cue of a predator. Furthermore, our results are the first to demonstrate that fish can acquire predator recognition through the pairing of a heterospecific alarm pheromone with the cue of a predator. These results suggest that brook sticklebacks will benefit by being in close proximity to fathead minnows. Acquired predator recognition has long-term consequences in mediating predator-prey interactions.     

Chemical Recognition of Risky Habitats is Culturally Transmitted among Fathead Minnows,Pimephales promelas (Osteichthyes,Cyprinidae)

Fathead minnows (Pimephales promelas) have an alarm substance (AS), or 'Schreckstoff', in epidermal club cells. Mechanical damage to the skin, as caused by a predator attack, releases the AS. The area in which conspecifics detect AS may be considered dangerous or risky because of the high probability of a subsequent predator attack. We exposed fathead minnows to water from one of two habitats (an open-water site and a vegetated-cover site) that we mixed with either AS or a distilled water control. Upon subsequent exposure to water from these habitats alone, minnows showed an antipredator response to the water they experienced in conjunction with AS, but not to water they received in conjunction with the distilled water control. These results confirmed that minnows can be conditioned with AS to recognize chemical cues from high-risk habitats. Naive minnows present during the fright response of conditioned minnows also exhibited antipredator behaviour, and subsequently responded when tested alone. Our results demonstrate that learned recognition of high-risk habitats can be transmitted culturally, which may allow minnows to lower their risk of predation.     

Population differences in the reaction of minnows to alarm substance*

Experiments showed that minnows, Phoxinus phoxinus, sympatric with pike, Esox lucius, responded more vigorously to alarm substance than minnows from a population with no experience of pike predation in the wild. Minnows from the pike-sympatric (Dorset) population were more likely to hide and less likely to risk feeding than their pike-allopatric (Gwynedd) counterparts. The reaction to alarm substance in the pike-sympatric population was further increased when it was presented along with the visual stimulus of a ‘stalking’ model pike. When the Dorset minnows experienced both alarm substance and the pike model together they reduced their inspection behaviour to a level below that of the Gwynedd minnows. Minnows from the Gwynedd (pike-allopatric) population displayed increased levels of shoaling in the treatments in which alarm substance was used.     

Chemically mediated predator inspection behaviour in the absence of predator visual cues by a characin fish

Animals commonly approach (i.e. 'inspect') potential predators. Glowlight tetras, Hemigrammus erythrozonus, have previously been shown to inspect the combined chemical and visual cues originating from novel predators and to modify their inspection (approach) behaviour depending upon the predator's diet. We conducted two experiments to determine whether tetras would inspect the chemical cues of injured prey or the dietary cues of a novel predator in the absence of any visual cues. Shoals of glowlight tetras were exposed to either distilled water (control) or the skin extract of swordtail (lacking ostariophysan alarm pheromones) or the skin extract of tetra (with alarm pheromones). There was no significant difference in the frequency of predator inspection behaviour towards swordtail or tetra skin extract compared to the distilled water controls. In the second experiment, we exposed shoals of tetras to either distilled water or the odour of Jack Dempsey cichlids, Cichlasoma octofasciatum, which had been food deprived, or fed a diet of swordtails or tetras. There was no significant difference in the frequency of predator inspection behaviour towards the odour of the starved cichlids and the odour of the fed cichlids in either of the two diet treatments. However, when tetras were exposed to the odour of cichlids fed tetras, they took significantly longer to initiate an inspection visit, remained further from the source of the chemical cues and inspected in smaller groups, compared with the odour of a starved cichlid or a cichlid fed swordtails. These data strongly suggest that tetras will inspect chemical cues alone, but only if the cue contains information about the predator. Copyright 2000 The Association for the Study of Animal Behaviour.     

高体鳑鲏化学预警响应与“个性”行为的关联

化学通讯是水生动物最原始、最普遍和最主要的通讯方式之一。对捕食风险源的回避可以使猎物减少被捕食风险,但如果出现的化学信息并不代表真正的威胁,那么就会导致猎物减少觅食或求偶的机会,从而降低适合度。因此基于能量代价-生态收益的权衡可能导致动物行为决策与化学通讯模式的分化。"个性"行为（Personality）已被证实与动物的行为决定有关,然而,有关鱼类化学预警通讯与"个性"行为关联的研究至今鲜见报道。推测："个性"行为是鱼类的化学通讯模式多样化的重要内因。采捕了广泛分布于我国淡水水域的野生高体鳑鲏（Rhodeus ocellatus）并于半自然状态下探究了：（1）实验鱼对不同化学信息（池水对照组、柠檬新奇信息组、高浓度和低浓度化学预警信息（Chemical alarm cues,CAC）组）的行为响应,（2）实验鱼在新异环境（被转入新栖息地）、新异刺激（新异物理刺激）、新异食物资源下的"个性"行为及其与化学预警响应的关联。结果发现：（1）不同溶液化学信息对实验鱼静止时间与爆发游泳的变化以及摄食个体的比例等参数均有显著影响（P < 0.05）。其中,柠檬组（未知"假风险"）与对照组相比无显著差异（P > 0.05）,实验鱼对不同浓度CAC（已知"真风险"）表现出不同程度的行为响应,高浓度和低浓度CAC均导致摄食个体比例下降（P < 0.05）,但只有高浓度CAC导致静止时间和爆发游泳的变化增加（P < 0.05）。可见,实验鱼对外源化学信息有准确高效的行为应答。（2）实验鱼静止时间比对不同溶液化学预警响应的变化与新异刺激下的活跃性显著负相关（P < 0.05）,但与新异环境下的活跃性无关（P > 0.05）,提示"个性"行为与化学预警响应有关但这种关联可能又有一定的环境依赖性（例如测试环境的稳定性与可预测性）。     

Predator recognition learning in rainbow darters Etheostoma caeruleum: specific learning and neophobia

This study investigated whether rainbow darters Etheostoma caeruleum can learn to recognize unfamiliar predators through the process of classical conditioning. Etheostoma caeruleum were conditioned by exposing them simultaneously to their chemical alarm cues (a known fright stimulus) and either chemical cues from larval ringed salamanders Ambystoma annulatum (unfamiliar predator) or to a blank water cue (control). Conditioning could result in either specific learning of the A. annulatum cue or increased wariness in response to any novel cue (neophobia). To distinguish between these possibilities, E. caeruleum in both groups were exposed to either A. annulatum cues alone or to chemical cues from western rat snakes Pantherophis obsoletus (novel cue) 2 days after conditioning. Treatment (A. annulatum‐conditioned) E. caeruleum, but not control E. caeruleum, showed a fright response when they were exposed to both the conditioned (A. annulatum) and novel (P. obsoletus) cues, indicating increased sensitivity to new stimuli. When E. caeruleum were retested after an additional 32 days, however, the fright response occurred only following exposure to the conditioned (A. annulatum) stimulus, indicating that specific learning of the A. annulatum cue had been retained whereas the neophobia to novel stimuli was temporary.     

Influence of Body Size on the Responses of Fathead Minnows, Pimephales promelas, to Damselfly Alarm Cues

A wide diversity of aquatic organisms release chemical alarm cues upon encountering or being attacked by a predator. These alarm cues can be used by nearby individuals to assess local predation risk. Receivers warned by chemical alarm cues gain a survival benefit when encountering predators. Animals that are in the same prey guild (i.e. that co‐occur and share the same predators) may learn to recognize each others’ chemical alarm cues. This ability may confer an adaptive advantage if the prey animals are vulnerable to the same predators. However, if the prey grow to different sizes and as a consequence are no longer vulnerable to the same suite of predators, then there should no longer be an advantage for the prey to respond to each others’ alarm cues. In this study, we exposed small and large fathead minnows (Pimephales promelas) to cues from syntopic injured damselfly larvae (Enallagma boreale), cues from injured mealworm larvae (Tenebrio molitor) and to distilled water. Small minnows exhibited antipredatory behaviour and increased shelter use in response to injured damselfly cues but not to the controls of injured mealworm or distilled water. On the contrary, large minnows exhibited no significant change in shelter use in response to any of the injured cues. These data demonstrate that fathead minnows exhibit an antipredator response to damselfly alarm cues, but only when minnows are small and members of the same prey guild as damselfly larvae. These results demonstrate the considerable flexibility in the responses to heterospecific alarm cues.     

The role of experience and chemical alarm signalling in predator recognition by fathead minnows, Pimephales promelas

Young-of-the-year, predator-naive fathead minnows, Pimephales promelas , from a pikesympatric population did not respond to chemical stimuli from northern pike, Esox Indus , while wild-caught fish of the same age and size did. These results suggest that chemical predator recognition is a result of previous experience and not genetic factors, Wild young-of-the-year minnows responded to pike odour with a response intensity that was similar to that of older fish, demonstrating that the ability to recognize predators is learned within the first year. The intensity of response of wild minnows which had been maintained in a predator free environment for 1 year was similar to that of recently caught minnows of the same age, suggesting that reinforcement was not required for predator recognition to be retained. Naive minnows that were exposed simultaneously to chemical stimuli from pike (a neutral stimulus) and minnow alarm substance exhibited a fright response upon subsequent exposure to the pike stimulus alone. Predator-naive minnows exposed simultaneously to chemical stimuli from pike and glass-distilled water did not exhibit a fright response to the pike stimulus alone. These results demonstrate that fathead minnows can acquire predator recognition through releaserinduced recognition learning, thus confirming a known mechanism through which alarm substance may benefit the receivers of an alarm signal.     

Learned Recognition of Novel Predator Odour by Zebra Danios, Danio Rerio, Following Time-shifted Presentation of Alarm Cue And Predator Odour

Fishes in the superorder ostariophysi possess specialized epidermal cells that contain an alarm cue. Fish associate novel odours, such as the odour of a predator, with predation risk after a single, simultaneous exposure to the novel odour and alarm cue. Thereafter, the novel cue is recognized as an indicator of risk and its presence induces antipredator behaviour. Two common antipredator behaviours are reduction in activity and movement to the bottom. This phenomenon has been demonstrated many times in the laboratory setting for a variety of aquatic taxa. In nature however, the detection of novel predator odour may be time-shifted with respect to the detection of alarm cues. Is there a critical period immediately upon the detection of alarm cue in which associative learning can occur? We presented zebra danios, Danio rerio, with the odour of northern pike, Esox lucius, 5?min after presenting them with either alarm cue or water (control). During a predation event, 5?min is a long time. When later retested with pike odour alone, zebra fish conditioned with alarm cue significantly increased antipredator behaviour in terms of decreased activity and movement towards the bottom. Control fish did not recognize pike odour as dangerous when retested. These data show that learned recognition of predation risk is sufficiently robust to accommodate ecologically realistic temporal shifts in stimulus presentation.     

Learned Recognition of Heterospecific Alarm Signals: The Importance of a Mixed Predator Diet

A wide diversity of aquatic organisms release alarm signals upon being attacked by a predator. Alarm signals may 'warn' nearby individuals of danger. Moreover, the signals may be important in facilitating learned recognition of unknown stimuli. It is common for different prey species to respond to each other's chemical alarm signals. In many cases, the responses are learned but no learning mechanisms have been identified to date. In this study we tested whether prey fish can learn the identity of an unknown alarm signal when they detect it in association with conspecific alarm cues in the diet of a predator. Chemical alarm cues are known to be conserved in the diet of predators. We conditioned fathead minnows ( Pimephales promelas ) with chemical stimuli from predatory yellow perch ( Perca flavescens ) fed a mixed diet of minnows and brook stickleback ( Culaea inconstans ), perch fed a mixed diet of swordtails ( Xiphophorus helleri ) and stickleback or distilled water. Minnows were subsequently exposed to chemical alarm cues of injured stickleback alone. Those minnows previously conditioned with perch fed a mixed diet of minnows and stickleback increased their use of shelter and 'froze' significantly more than minnows previously conditioned with perch fed a diet of swordtails and stickleback or those exposed to distilled water. These data demonstrate a mechanism by which minnows can learn the identity of a heterospecific alarm signal.     

Sensory ecology of ostariophysan alarm substances

Chemical communication of predation risk has evolved multiple times in fish species, with conspecific alarm substance (CAS) being the most well understood mechanism. CAS is released after epithelial damage, usually when prey fish are captured by a predator and elicits neurobehavioural adjustments in conspecifics which increase the probability of avoiding predation. As such, CAS is a partial predator stimulus, eliciting risk assessment-like and avoidance behaviours and disrupting the predation sequence. The present paper reviews the distribution and putative composition of CAS in fish and presents a model for the neural processing of these structures by the olfactory and the brain aversive systems. Applications of CAS in the behavioural neurosciences and neuropharmacology are also presented, exploiting the potential of model fish [e.g., zebrafish Danio rerio, guppies Poecilia reticulata, minnows Phoxinus phoxinus) in neurobehavioural research.     

Detection of conspecific alarm cues by juvenile salmonids under neutral and weakly acidic conditions: laboratory and field tests

A variety of fishes possess damage-released chemical alarm cues, which play a critical role in the detection and avoidance of potential predation threats. Recently, we have demonstrated that the ability of fathead minnows (Pimephales promelas) and finescale dace (Phoxinus neogaeus) to detect and respond to conspecific alarm cues is significantly reduced under weakly acidic conditions (pH 6.0). Rainbow trout (Oncorhynchus mykiss) and brook charr (Salvelinus fontinalis) possess an analogous alarm cue system. However, it is unknown if the trout alarm cue system is likewise affected by relatively small changes in pH. In addition, previous studies have not verified this phenomenon under natural conditions. We conducted laboratory and field trials to examine the potential effects of acute exposure to weakly acidic (pH 6.0) conditions on the detection and response of conspecific alarm cues by juvenile trout. Our laboratory results demonstrate that while juvenile rainbow trout exhibit significant increases in antipredator behaviour under normal pH conditions (pH 7.0–7.2), they do not respond to the presence of conspecific chemical alarm cues (i.e. response is not different from controls) under weakly acidic conditions. Similarly, a wild strain of brook charr in their natural streams near Sudbury, Ontario, failed to detect conspecific alarm cues in a weakly acidic stream (mean pH 6.11) while they responded to these cues in a neutral stream (mean pH of 6.88). This is the first demonstration that relatively small changes in ambient pH can influence alarm responses under natural conditions. These data suggest significant, sub-lethal effects of acid precipitation on natural waterways.     

Antipredator responses to skin extract of redbelly dace, <Emphasis Type="Italic">Phoxinus eos, </Emphasis>by free-ranging populations of redbelly dace and fathead minnows,<Emphasis Type="Italic">Pimephales promelas</Emphasis>

Synopsis Fishes in the superorder Ostariophysi possess specialized epidermal cells that contain a chemical alarm cue. The alarm cue is released when the skin is damaged during a predatory attack. Therefore, the cue serves as a reliable indicator of predation risk to nearby conspecifics and ecologically similar heterospecifics with which it shares predators. Antipredator behavior in response to these alarm cues has been demonstrated in numerous studies in confined spaces (laboratory aquaria, field traps, fluvarium). When tested on a natural field population however, behavioral response has been inconsistent. Here, we expose free-ranging redbelly dace and fathead minnows to skin extract of redbelly dace and record their behavioral response with an underwater video camera. We observed avoidance of areas in which skin extract was introduced, but no avoidance of areas in which water (control) was introduced. These data confirm the ecological function of skin extract in mediating predator–prey interactions in aquatic habitats, and argue against the hypothesis that alarm reactions are an artifact of confined spaces.