The Significance of Antecedent Auditory Experiences on Later Reactions to the ‘Seeet’ Alarm-Call in Great Tit Nestlings Parus major

Like many other naive young birds, great tit nestlings, Parus major, respond by a persistent cessation-of-begging and immobility response to the ‘seeet’ alarm-call of the species (Rydén 1978). The experimental antecedents of this reaction were investigated through a systematic manipulation of the auditory environment preceding it. Limiting nestlings' experience with the ‘seeet’-call to either a “positive”, a “neutral” or a “negative” setting led to their normal aversive response to the call being weakened, unaltered and strengthened, respectively. Nestlings brought up in a “reversed”, artificial environment showed a radical decrease in their aversive response. The results lend support to the hypotheses that a) the ‘seeet’-call owes its aversive effect to the fact that its acoustic character contrasts sharply with the main features of the nestlings' normal auditory environment; b) context-specific experiences may either increase or decrease this basic response tendency. Some functional implications of the results as related to the behavior of the species in nature are discussed.     

Two conflicting needs affecting predator mobbing by great tits,Parus major

Predator mobbing by great tits, Parus major, was studied by manipulating two contextual variables: the proximity of danger to the nest hole (close versus far), and breeding condition (non-breeding versus nest building versus care of nestlings). In five experiments pitting these contextual variables against each other, great tits were presented with a multi-species mobbing chorus signalling danger to which they responded by typical predator harassment. Based on the ‘move on’ hypothesis of harassment and on context manipulation, harassment levels specific for each of the five experiments could be predictably ordered; the ensuing rank order of scores reflected, by its nature, a series of four strong inference tests. Whereas two latency measures supported the move on hypothesis, two distance-from-danger measures suggested nest hole concealment to be an opposing need; a fifth measure of harassment, comprising both latency and distance, held an intermediate position, thus tying in with the compromise nature of harassment. Considering the ‘phantom’ nature of the unseen danger stimulus applied, the defence levels observed must be regarded as moderately conservative. There are natural danger stimuli (e.g. cats) that more effectively prevent great tits displaying predator harassment.     

The Feeding Behaviour of Starlings (Sturnus vulgaris) in the Presence of ‘Eyes’

Previous studies that have looked at the aversive properties of simple, predator-related, eye-like patterns presented in an artificial context, have yielded conflicting results. The aim of the present series of experiments was to investigate whether such stimuli had potential for use as bird scarers. The starling (Sturnus vulgaris) was the subject species. The experimental procedure was designed to provide a relevant and objective, interval scale of aversiveness based upon the ability of eye patterns to deter hungry birds from a feeding area. Each subject was deprived of food for 5 h and then tested in an apparatus which exposed an eye pattern over a food trough immediately after the bird alighted on that trough. 10 measures of the starlings' feeding behaviour were monitored over a 1-h trial. It was predicted that five of these variables would be positively and five negatively correlated with an increase in the fear evoking properties of the stimulus. The 10 measures were analysed using principal component analysis and the first component in every experiment had the signs of the latent vectors of the variables exactly in the predicted pattern and therefore appeared to be an ‘aversiveness index’. The scores from this first component were then used in a Latin square ANOVA to distinguish subject, test order and stimulus effects. The main findings from the eight experiments are as follows. Simple eye-like patterns can indeed deter hungry starlings from feeding in their vicinity. The presence of a pair of eyes painted on a white card caused an 88% reduction in time spent on the nearby food trough and a 65% reduction in actual feeding time. The presence of ‘pupils’ in the patterns is essential for simple circles alone are not significantly more aversive than the control card. Once a pattern has the ‘pupil/iris’ distinction then a circular outline becomes important although the shape of the ‘pupil’ appears relatively unimportant. Eyes with coloured irises are more effective than black and white patterns but the degree of contrast between the ‘pupil’ and ‘iris’ appears irrelevant. Increasing the number of simultaneously presented eyes from one to three is correlated with an increasing trend in aversiveness which is destroyed if the same stimuli are presented within the context of a simple head outline. Changing the orientation of a pair of eyes from horizontal to vertical only slightly reduces their aversiveness. The size of the patterns appears to be unimportant within the range tested. The aversive properties of eyespots and of broadcast starling distress calls are positively additive. The effectiveness of a stimulus combining all the important features mentioned is as aversive as is a realistic model of the head of an owl.     

THE ABILITY OF BARN OWLS TYTO ALBA TO DISCRIMINATE AND LOCALIZE AVIAN ALARM CALLS

Ten Barn Owls were tested in a sound-room with recordings of various alarm calls and noise. The owls readily responded with head movements to the majority of stimuli and correctly oriented to their source. The seeet alarm call and the 'warning' call elicited the least responses, but they were properly localized by all owls which did respond. This suggests that rather than non-localizability accounting for the fewer observed responses to the seeet and 'warning' calls, there is either something else in the quality of those sounds, or in the manner in which they are processed by the bird, which does not elicit responses to the degree that transient sounds with broad frequencies do. Selection in these predators for non-responsiveness is suggested as a counter-strategy to minimize futile energy expenditure in attacking, or otherwise responding to an alerted prey.
An additional hypothesis, consistent with Darwin's antithesis principle, is proposed whereby intraspecific and interspecific selection, rather than predator selection for non-localizability, has led to the evolution of seeet and mobbing calls.     

Sex and age differences in reflectance and biochemistry of carotenoid‐based colour variation in the great tit Parus major

The plumage coloration in great tits (Parus major) is the subject of much behavioural and ecophysiological research, yet there is a lack of analyses of the natural colour variation and its mechanisms. We used reflectance spectrometry and high‐performance liquid chromatography to explore individual, sexual and age‐related variation in carotenoid coloration and pigmentation, paramount to the often presumed, but rarely substantiated, costs and ‘honesty’ of carotenoid displays. In adults, we found that sex was the strongest predictor of ‘brightness’ (higher in males) and of ‘hue’ (longer wavelength in females). There was no sex difference in ‘carotenoid chroma’ or carotenoid content of feathers which also was unrelated to adult age (1 or 2+ years) and condition. Similar patterns were revealed for nestlings. Regarding the biochemical ‘signal content’, ‘carotenoid chroma’, but not ‘hue’, was significantly related to the carotenoid content (lutein and zeaxanthin) of feathers. These results refute the previously assumed exaggeration of carotenoid pigmentation in male great tits, and question the condition‐dependence of carotenoid coloration in this species. However, the sexual dimorphism in total reflectance or ‘brightness’, most likely due to melanins rather than carotenoids, may have implications for signalling or other adaptive explanations that need to be explored. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 758–765.     

Horned larks on the Tibetan Plateau adjust the breeding strategy according to the seasonal changes in the risk of nest predation and food availability

Songbirds in seasonal environments often adjust their breeding strategy according to spatial or temporal changes in breeding conditions. Here we investigate how horned larks Eremophila alpestris, a multi‐brooded songbird on the Tibetan Plateau, responded to the changing risk of nest predation and food availability across breeding attempts. We showed that both nest concealment and food supply increased with plant growth, and horned larks adjusted their breeding strategies accordingly. First they selected nest‐sites where predator density was low, which enhanced nest survival. Second, clutch size increased with improving breeding conditions. They did not adopt an ‘egg‐size’ strategy as egg size did not change with laying sequence or breeding attempt. Instead, they adopted the ‘brood survival (feeding later‐hatched nestlings more)’ and ‘brood reduction (feeding early‐hatched nestlings more)’ strategies during early and later attempts. Moreover, nestlings’ growth varied with breeding attempt: more energy was invested into the growth of body mass during the first attempt but more energy was expended on the growth of linear structures during later attempts. This difference in energy allocation reflected changing food availability. We suggest that temporal changes of environmental factors are also the important force driving the evolution of avian breeding strategies.     

Parental investment in nest defence by stonechats (Saxicola torquata)

Breeding stonechats (Saxicola torquata) made mixed sequences of two calls when a human intruder entered territories. ‘Whits’ are modulated notes with a small frequency range, and in laboratory tests caused nestlings to stop begging. ‘Chacks’ cover a wide range of frequencies, and in the field were combined with flights made so as to distract an intruder from the nest. On average male and female call-rates were similar, but varied greatly according to the intruder's distance from the nest, and at different stages of the nesting cycle. Rates increased rapidly after hatching, and this correlated most closely with the cumulative total of parents' visits to feed nestlings. This suggests that the level of defence may be adjusted to the value of the offspring to their parents. Call-rates declined about one week after fledging. A smaller peak by some pairs at the start of incubation was apparently related to probable poor condition after a previous breeding attempt, and after laying large clutches. Rates of Whits were higher at nests with larger broods, up to an asymptote, but rates of Chacks were independent of brood size. Birds suffering nest-predation showed lower call-rates before the event than equivalent successful birds, suggesting that the calls do reduce the risk of predation.     

Pain as a Cause of Aggression

Aggressive behavior can be elicited by aversive stimuli as anunconditioned reflex. Electric shock, heat, physical blows,and intra-cranial stimulation are among the stimuli which willelicit fighting. Various characteristics of the stimulus—suchas frequency of presentation, intensity, and duration, in thecase of shock—and of the environment in general, influencethe rate at which pain-elicited fighting occurs. Characteristicsof the animals—such as sex, age, sensory impairment, species,and social experience— also influence the occurrence ofrespondent aggression. Aggression can be conditioned according to both the operantand the respondent paradigms. Since both operant and respondentaggression can occur concurrently, interesting interactionsresult. Depending upon a number of variables, respondent aggressionexerts a disruptive influence on social behavior controlledby aversive stimuli. The interference of aggression with behaviormaintained by aversive stimuli may cause us to revise our opinionof the efficiency of aversive stimuli in social situations. Although pain is probably a source of human aggression, furtherresearch with infra-human animals, as well as actual researchwith humans, should elucidate the relationship between aversivestimuli and aggression in man.     

Relationship between Fear Conditionability and Aversive Memories: Evidence from a Novel Conditioned-Intrusion Paradigm

Intrusive memories – a hallmark symptom of posttraumatic stress disorder (PTSD) – are often triggered by stimuli possessing similarity with cues that predicted or accompanied the traumatic event. According to learning theories, intrusive memories can be seen as a conditioned response to trauma reminders. However, direct laboratory evidence for the link between fear conditionability and intrusive memories is missing. Furthermore, fear conditioning studies have predominantly relied on standardized aversive stimuli (e.g. electric stimulation) that bear little resemblance to typical traumatic events. To investigate the general relationship between fear conditionability and aversive memories, we tested 66 mentally healthy females in a novel conditioned-intrusion paradigm designed to model real-life traumatic experiences. The paradigm included a differential fear conditioning procedure with neutral sounds as conditioned stimuli and short violent film clips as unconditioned stimuli. Subsequent aversive memories were assessed through a memory triggering task (within 30 minutes, in the laboratory) and ambulatory assessment (involuntary aversive memories in the 2 days following the experiment). Skin conductance responses and subjective ratings demonstrated successful differential conditioning indicating that naturalistic aversive film stimuli can be used in a fear conditioning experiment. Furthermore, aversive memories were elicited in response to the conditioned stimuli during the memory triggering task and also occurred in the 2 days following the experiment. Importantly, participants who displayed higher conditionability showed more aversive memories during the memory triggering task and during ambulatory assessment. This suggests that fear conditioning constitutes an important source of persistent aversive memories. Implications for PTSD and its treatment are discussed.     

Effects of predictability on the welfare of captive animals

Variations in the predictability of a stressor have pronounced effects on the behavioural and physiological effects of stress in rats. It is reasonable to expect that variations in the predictability of husbandry routines thought to be aversive to animals might have similar effects on stress indices. Similarly, variations in the predictability of positive events, of which feeding is an obvious example, may affect welfare. This review examines the behavioural and physiological effects of the predictability of aversive and appetitive stimuli, and the application of experimental findings to animal husbandry in practice. It is argued here that two distinct but overlapping types of predictability exist. ‘Temporal’ predictability describes whether an event occurs at fixed or variable intervals, whereas ‘signalled’ predictability relates to the reliability of a signal preceding the event. This review examines the effects of each of these types of predictability in relation to positively and negatively perceived events, and examines the link between predictability and control. Recommendations are made for relatively simple and inexpensive modifications to husbandry routines that may be easy to incorporate into the schedules of busy staff yet could have a profound impact on the welfare of animals in their care.     

Training methods of military dog handlers and their effects on the team's performances

While only a few studies have analysed training methods used on working dogs, a recent survey in 303 Belgian military handlers revealed the use of harsh training methods on military working dogs (MWD). The present work aims at analysing the training methods used on Belgian MWD and the behaviour of handlers to objectify the performances of the dog handlers teams (DH teams) and the welfare of the animals.A standardized evaluation, including obedience and protection work exercises, was conducted on DH teams (n = 33). Every evaluation was done twice to assess the reliability of the observation methods. The behaviours of MWD and handlers were recorded on videotape and subsequently analysed. Results showed that handlers rewarded or punished their dogs intermittently. Stroking and patting the dogs were the most frequently used rewards. Pulling on the leash and hanging dogs by their collars were the most commonly used aversive stimuli.The team's performance was influenced by the training method and by the dog's concentration: (1) low-performance dogs received more aversive stimuli than high-performance dogs; (2) dog's distraction influenced the performance: distracted dogs performed less well.Handlers punished more and rewarded less at the second evaluation than at the first one. This suggests that handlers modified their usual behaviour at the first evaluation in view to present themselves in a positive light. During the second evaluation the dogs reacted to this higher frequency of aversive stimuli as they exhibited a lower posture after aversive stimuli. The authors cannot prove that the welfare of these dogs had been hampered, but there is an indication that it was under threat.Low team performances suggest that DH teams should train more regularly and undertake the usefulness of setting a new training system that would rely on: the use of more positive training methods, an increased training frequency, the elaboration of a course on training principles, and an improvement of dog handler relationship.     

Wing-flick Signals in the Courtship of the African Cave Cricket,Phaeophilacris spectrum

The male of the African cave cricket Phaeophilacris spectrum (Saltatoria: Grylloidea: Phalangopsidae) possesses tegmina without stridulatory organs, and both females and males lack tympanal organs. Therefore acoustic communication in the usual sense, which is typical for most crickets, is absent in this species. However, adaptations of the wing articulation allow the males to flick their wings forward over their heads. During courtship these movements are performed in series of 4–5 wing-flicks at a rate of 8–12 Hz (called wing-flick series = WFS) which elicit low-frequency air movements. Such signals may have taken over the functional significance of an acoustic ‘courtship song’. A quantitative analysis of the cricket's behaviour showed that in terms of frequency and total duration, ‘wing-flick series’ and ‘rocking’ behaviour are the most prominent signals of the male's courtship display. Moreover, analysis of the spatial relationships between male and female demonstrates that the wing-flicking is directed towards the female. The response of a female to a wing-flicking, courting male seems mainly to be calmed down and to reach a passive, receptive state, necessary for subsequent copulation. Females confronted with a male's courtship display exhibited a significantly higher threshold to disturbing stimuli than uncourted ones.     

‘Communication’ in weakly electric fish, Gnathonemus niger (Mormyridae) I. Variation of electric organ discharge (EOD) frequency elicited by controlled electric stimuli

The weakly electric fish, Gathonemus niger, discharged with a frequency of 4 to 8 Hz during the day and 10 to 16 Hz during the night. The frequency of superimposed burst discharges (32 to 56 Hz) was independent of diurnal factors. The variation of the electric organ discharge frequency during the day was investigated in response to controlled electric stimulus patterns: (a) A free running stimulus frequency of 4 Hz, simulating the resting frequency of another fish, and different stimulus intensities, simulating different distances between two fish. (b) Free running frequencies of 4, 8, 16, …, 128 Hz and two particular stimulus intensities. (c) Discharge coupled stimuli (each discharge triggered an electric stimulus with a fixed delay) and different stimulus intensities.All three kinds of stimuli elicited defined and predictable response discharge patterns supporting the assumption that an electric fish would respond to a particular discharge pattern of another fish also in a similar and predictable manner. Low stimulus intensities (0·04 to 0·2 mV per cm) caused cessation of the discharge activity, a ‘hiding’ or ‘listening’ response. The discharge rate increased linearly with the logarithm of the stimulus intensity. The fish was particularly sensitive to stimulus frequencies which simulated its burst activity (32 to 56 Hz). Discharge coupled stimuli showed that the fish responded to about eight times lower stimulus intensities if the stimulus occurred between two discharges (15 to 30 m-s after the fish's discharge) than if the stimulus occurred within or immediately after the discharge. All suprathreshold stimuli elicited a typical discharge pattern: The irregular resting discharge activity became significantly regular. The degree of regularity was even improved during maintained stimulation. The regularisation of the discharge activity is thought to be involved in the fish's electrolocating system whereas frequency variations are considered as being involved in both the locating system and as communication signals among weakly electric fish.     

P3 responses to prosodic stimuli in adult autistic subjects

Autistic persons are known to have serious abnormalities in speech prosody. The present study attempted to ascertain whether autistic persons could discriminate and/or recognize prosodic contrasts in auditory stimuli. A group of 11 adult autistic subjects with normal IQ and an age-matched group of normal subjects were studied electrophysiologically and behaviorally during presentations of prosodic and phonemic stimuli. The cognitive P3 potential was recorded in response to rare (20%)/frequent (80%) presentations of phonemic stimuli, ‘ba/pa’, linguistic-prosodic stimuli, ‘Bob’. (statement)/‘Bob?’ (question), and emotional-prosodic stimuli, ‘Bob’ (happy)/‘Bob’ (happy)/’Bob’ (angry). Behaviorally, auditory discrimination was tested by requiring a button-press response to each presentation of the rare target stimulus and cognitive association was tested by requiring a match between the verbalized stimulus and an appropriate picture/word.Contrary to our hypothesis, the autistic subjects generally showed normal P3 responses to all stimuli and performed at a normal level in all behavioral tests. However, a significant autistic P3 response to the phoneme ‘pa’ was not demonstrated. This surprising result was reexamined and shown to reflect an unusually large autistic response to ‘pa’ as the frequent stimulus in the first recording block, this initial hyper-reactivity prevented a ‘frequent/rare’ differential when ‘pa’ was presented as the rare stimulus in a later recording block. In the P3 latency window, both the autistic and control groups showed the largest amplitude responses to emotional-prosodic stimuli; neither the N1 nor P2 showed these stimulus effects. Thus, ‘emotional sounds’ appear to be particularly effective in activating the neural substrate of the P3 generator system.Overall, these data indicate remarkably normal P3 and behavioral processing of prosodic stimuli by the high-functioning autistic subjects of this study.     

Timing of breeding and nestling diet of Wood Warbler Phylloscopus sibilatrix in relation to changing food supply

Capsule Wood Warblers did not match their reproduction to the caterpillar peak

Aims To study the timing of Wood Warblers’ breeding and nestling diet in relation to caterpillar abundance under primeval conditions in the Bia?owie?a National Park, Poland.

Methods Observations of food brought by parents. Inter‐year and seasonal changes in availability of folivorous caterpillars were assessed by direct counts and caterpillar frass collection.

Results Maximum food requirements occurred two weeks after the peak of caterpillar abundance. The mismatch had no effect on nestlings’ development. Diet varied little across years and habitats, but varied strongly within a season. Following the decline of ‘green’ caterpillars, their proportion in the Wood Warbler diet strongly declined. Caterpillars were replaced by winged insects. Small nestlings received more spiders than older ones.

Conclusion Timing of Wood Warbler breeding in Bia?owie?a National Park was constrained by the females’ arrival time. Birds did not match their reproduction to the caterpillar peak. ‘Green’ caterpillars were the preferred food for nestlings; birds responded to the caterpillar decrease by feeding smaller proportions of them and switching to alternative prey, namely winged insects.     

Habitat fragmentation influences nestling growth in Mediterranean blue and great tits

In patchy forest areas, the size of the forest patch where birds breed has a strong influence on their breeding success. However, the proximate effects contributing to lowering the breeding success in small forest patches remain unclear; and a shortage of crucial resources in those forest patches has been suggested to account in some degree for this failure. With the aim to further investigate this issue, we have monitored the breeding cycle of blue and great tits in three ‘large’ forest patches (ranging between 26.5 and 29.6 ha) and twelve ‘small’ forest patches (ranging between 1.1 and 2.1 ha) in a Mediterranean area in central Spain, during three years (2011–2013). We also recorded the nestling diet inside the nest-boxes with the aid of handy-cams. Only males significantly differed between forest patch size categories; being on average younger and with better body condition in small patches for great and blue tits respectively. Reproductive traits did not vary between forest patch size categories, but the body condition of blue tit nestlings and the size of great tit nestlings did, being significantly better and larger respectively in large forest patches. The recruitment rate of blue tit nestlings was also higher in large patches. Regarding nestling diet, blue tits did not differ but great tits did, delivering a larger amount of caterpillars in large forest patches. Most variation in the reproductive traits occurred between years, probably due to annual differences in environmental conditions. This study suggests that food supply could be limiting the breeding success of birds above all in small patches, but also in large patches under particular environmental conditions.     

Information processing in a cricket ganglion: The response of giant fibres to sound pulses

The response of giant fibres in the ventral nerve cord to stimulation of cercal afferents with pulses of sound was studied in the domestic cricket, Acheta domesticus. Pulses at 450 Hz gave the highest frequency response in several classes of units, and were therefore used as stimuli in subsequent experiments. In intact animals the response of the giant fibres to bilateral cercal stimulation showed a characteristic high frequency ‘on’ response followed by steady firing of some units for the duration of the sound pulse. The end of each pulse was followed by a short period of inhibition of the tonic units.Cercal amputation and other experiments showed that input from cercal afferents excites both large and small ipsilateral giants, and excites small and inhibits large contralateral giants. Descending input from higher neural centres in intact animals tends to reduce the responses to the stimuli. It is suggested that a function of the contralateral excitatory and inhibitory effects is to sharpen the ‘on’ response of the giant fibres to sound stimuli in intact animals.     

Frustration Behaviors in Domestic Dogs

During extinction a previously learned behavior stops being reinforced. In addition to the decrease in the rate of the instrumental response, it produces an aversive emotional state known as frustration. This state can be assimilated with the fear reactions that occur after aversive stimuli are introduced at both the physiological and behavioral levels. This study evaluated frustration reactions of domestic dogs (Canis familiaris) during a communicative situation involving interactions with a human. The task included the reinforcement and extinction of the gaze response toward the experimenter's face when the dogs tried to obtain inaccessible food. The dog's frustration reactions during extinction involved an increase in withdrawal and side orientation to the location of the human as well as lying down, ambulation, sniffing, and vocalizations compared with the last acquisition trial. These results are especially relevant for domestic dog training situations in which the extinction technique is commonly used to discourage undesirable behaviors.     

Nucleus accumbens neurons are innately tuned for rewarding and aversive taste stimuli, encode their predictors, and are linked to motor output

The nucleus accumbens (NAc) is a key component of the brain's reward pathway, yet little is known of how NAc cells respond to primary rewarding or aversive stimuli. Here, naive rats received brief intraoral infusions of sucrose and quinine paired with cues in a classical conditioning paradigm while the electrophysiological activity of individual NAc neurons was recorded. NAc neurons (102) were typically inhibited by sucrose (39 of 52, 75%) or excited by quinine (30 of 40, 75%) infusions. Changes in firing rate were correlated with the oromotor response to intraoral infusions. Most taste-responsive neurons responded to only one of the stimuli. NAc neurons developed responses to the cues paired with sucrose and quinine. Thus, NAc neurons are innately tuned to rewarding and aversive stimuli and rapidly develop responses to predictive cues. The results indicate that the output of the NAc is very different when rats taste rewarding versus aversive stimuli.