Sexual-discrimination learning in male Japanese quail (Coturnix coturnix japonica)

We investigated how male Japanese quail (Coturnix coturnix japonica) learn through extensive social and sexual experience to discriminate between male and female conspecifics. Opportunity for heterosexual copulation was important for this learning, but even extensive copulatory opportunity was not sufficient to produce a sexual discrimination; subjects also required exposure to other males. Exposure to females after copulatory opportunity did not produce a sexual discrimination but facilitated its acquisition. Time or exposure to only the visual features of male birds (provided by taxidermic models) after copulatory opportunity did not result in differential responding to male and female conspecifics. Finally, presenting stimulus birds one at a time proved to be a more sensitive test of sexual-discrimination learning than presenting two stimulus birds at the same time. The results indicate that sexual-discrimination learning is similar to conventional associative learning.     

Thinking chickens: a review of cognition,emotion, and behavior in the domestic chicken

Domestic chickens are members of an order, Aves, which has been the focus of a revolution in our understanding of neuroanatomical, cognitive, and social complexity. At least some birds are now known to be on par with many mammals in terms of their level of intelligence, emotional sophistication, and social interaction. Yet, views of chickens have largely remained unrevised by this new evidence. In this paper, I examine the peer-reviewed scientific data on the leading edge of cognition, emotions, personality, and sociality in chickens, exploring such areas as self-awareness, cognitive bias, social learning and self-control, and comparing their abilities in these areas with other birds and other vertebrates, particularly mammals. My overall conclusion is that chickens are just as cognitively, emotionally and socially complex as most other birds and mammals in many areas, and that there is a need for further noninvasive comparative behavioral research with chickens as well as a re-framing of current views about their intelligence.     

Worms under cover: relationships between performance in learning tasks and personality in great tits (Parus major)

In animals, individual differences in learning ability are common and are in part explained by genetic differences, developmental conditions and by general experience. Yet, not all variations in learning are well understood. Individual differences in learning may be associated with elementary individual characteristics that are consistent across situations and over time, commonly referred to as personality or temperament. Here, we tested whether or not male great tits (Parus major) from two selection lines for fast or slow exploratory behaviour, an operational measure for avian personality, vary in their learning performance in two related consecutive tasks. In the first task, birds had to associate a colour with a reward whereas in the second task, they had to associate a new colour with a reward ignoring the previously rewarded colour. Slow explorers had shorter latencies to approach the experimental device compared with fast explorers in both tasks, but birds from the two selection lines did not differ in accomplishing the first task, that is, to associate a colour with a reward. However, in the second task, fast explorers had longer latencies to solve the trials than slow explorers. Moreover, relative to the number of trials needed to reach the learning criteria in the first task, birds from the slow selection line took more trials to associate a new colour with a reward while ignoring the previously learned association compared with birds from the fast selection line. Overall, the experiments suggest that personality in great tits is not strongly related to learning per se in such an association task, but that birds from different selection lines might express different learning strategies as birds from the different selection lines were differently affected by their previous learning performance.     

Blockade of NMDA Receptors in the Anterior Forebrain Impairs Sensory Acquisition in the Zebra Finch (Poephila guttata)

Juvenile zebra finches (Poephila guttata) learn song in two stages: during sensory acquisition, they memorize the song of an adult tutor, and during sensorimotor learning, they alter their vocalizations to match the stored song model. Like many other forms of neural plasticity and memory formation, vocal learning in zebra finches is impaired by pharmacological blockade of NMDA receptors, but the relevant NMDA receptors have not yet been localized. During song development, one neural region that has been implicated specifically in song learning, the lMAN, exhibits an increased density of NMDA receptors as well as decreased binding affinity for the NMDA antagonist MK-801. To test the hypothesis that sensory acquisition requires activation of NMDA receptors in or near the lMAN we infused the NMDA receptor antagonist amino-5-phosphonopentanoic acid (AP5; 2.5 μg in 0.1 μl) directly into the anterior forebrain. Birds receiving AP5 infusions prior to each of 10 tutoring sessions copied significantly less of their tutor's song than did sham-operated birds, saline-infused birds, birds that received AP5 infusions on nontutoring days, or birds that received AP5 infusions into the cerebellum. Furthermore, infusions of AP5 in the anterior forebrain did not impair young birds’ ability to discriminate zebra finch from canary song. These findings are consistent with the hypothesis that NMDA receptor activation in the anterior forebrain is necessary for the memorization of song material during avian vocal learning. This is also the first report that song-related regions of the anterior forebrain contribute to sensory acquisition specifically.     

The effect of social learning on avoidance of aposematic prey in juvenile great tits (<Emphasis Type="Italic">Parus major</Emphasis>)

Social learning plays an important role in acquiring new foraging skills and food preferences in many bird species but its potential role in learning to avoid aposematic prey has never been studied. We tested the effect of social learning on the acquisition of avoidance of aposematic insect prey (firebug Pyrrhocoris apterus; Heteroptera) in juvenile, hand-reared great tits (Parus major). Behaviour towards aposematic prey was compared between two groups of birds: (1) the observers that were, prior to encounter with firebugs, allowed to watch the experienced conspecific demonstrator repeatedly refuse to attack the prey, and (2) the control birds that lacked this opportunity. Observing an experienced demonstrator was not sufficient for learning complete avoidance, because birds from both groups attacked at least the first firebug they had encountered in avoidance training. However, the opportunity to observe the avoidance behaviour of another bird significantly increased the rate of subsequent individual learning of observers in comparison with control birds. Social learning also decreased mortality of firebugs killed by the birds during the avoidance learning. Socially enhanced learning to avoid aposematic prey might be a mechanism important especially for naive juvenile birds learning from their parents, but it could also enhance learning in adults from their more experienced flock mates. Because social learning of avoidance may also lead to decreased mortality of aposematic prey, its effect should be taken into account in scenarios considering evolution and maintenance of prey warning signals.     

Selective impairment of song learning following lesions of a forebrain nucleus in the juvenile zebra finch

Area X, a large sexually dimorphic nucleus in the avian ventral forebrain, is part of a highly discrete system of interconnected nuclei that have been implicated in either song learning or adult song production. Previously, this nucleus has been included in the song system because of its substantial connections with other vocal control nuclei, and because its volume is positively correlated with the capacity for song. In order to directly assess the role of Area X in song behavior, this nucleus was bilaterally lesioned in both juvenile and adult zebra finches, using ibotenic acid. We report here that lesioning Area X disrupts normal song development in juvenile birds, but does not affect the production of stereotyped song by adult birds. Although juvenile-lesioned birds were consistently judged as being in earlier stages of vocal development than age-matched controls, they continued to produce normal song-like vocalizations. Thus, unlike the lateral magnocellular nucleus of the anterior neostriatum, another avian forebrain nucleus implicated in song learning, Area X does not seem to be necessary for sustaining production of juvenile song. Rather, the behavioral results suggest Area X is important for either the acquisition of a song model or the improvement of song through vocal practice.     

Attention control: Relationships between self-report and behavioural measures,and symptoms of anxiety and depression

Previous research has demonstrated differences in processing between fear-relevant stimuli, such as snakes and spiders, and non-fear-relevant stimuli. The current research examined whether non-fear-relevant animal stimuli, such as dogs, birds and fish, were processed like fear-relevant stimuli following aversive learning. Pictures of a priori fear-relevant animals, snakes and spiders, were evaluated as negative in affective priming and ratings and were preferentially attended to in a visual search task. Pictures of dogs, birds and fish that had been trained as CS+ in an aversive conditioning design were evaluated more negatively and facilitated dot probe detection relative to CS? pictures. The current studies demonstrated that stimuli viewed as positive prior to aversive learning were negative and were preferentially attended to after a brief learning episode. We propose that aversive learning may provide a mechanism for the acquisition of stimulus fear relevance.     

Matching and oddity relational learning by pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>): transfer from color to shape

Relational learning, as opposed to perceptual learning, is based on the abstract properties of the stimuli. Although at present there is no doubt that pigeons are capable of relational behavior, this study aims to further disclose the conditions under which it occurs. Pigeons were trained in an outdoor cage on a matching-to-sample or an oddity-from-sample task, with colored cardboard stimuli presented horizontally. The apparatus involved three sliding lids on which the stimuli were drawn and which, when displaced, revealed the reinforcement. The lids were either adjacent to each other or somewhat separated. Training sessions involved two colors, and test sessions six different colors (same dimension test), or six different shapes (different dimension test). One group of birds trained under the ‘adjacent’ condition failed when tested with new stimuli, but succeeded in both dimension tests after training under the ‘separate’ condition. Two other groups of birds succeeded in all tests after training under the latter condition. These results show that depending on procedural details, pigeons are or are not able to transfer from one visual dimension to another, thus extending previous related findings.     

Direct social contacts override auditory information in the song-learning process in starlings (Sturnus vulgaris)

Social influence on song acquisition was studied in 3 groups of young European starlings raised under different social conditions but with the same auditory experience of adult song. Attentional focusing on preferred partners appears the most likely explanation for differences found in song acquisition in relation to experience, sex, and song categories. Thus, pair-isolated birds learned from each other and not from broadcast live songs, females did not learn from the adult male tutors, and sharing occurred more between socially associated peers. On the contrary, single-isolated birds clearly copied the adult songs that may have been the only source of attention stimulation. Therefore, social preference appears as both a motor for song learning and a potential obstacle for acquisition from nonpreferred partners, including adults.     

Discriminating the sex of conspecifics by male Japanese quail (Coturnix coturnix japonica).

How male Japanese quail (Coturnix coturnix japonica) discriminate the sex of conspecifics at a distance was investigated by testing subjects with male and female stimulus birds on the other side of a small window after opportunities to copulate with females and again after repeated exposures to males in the absence of copulatory opportunity. Time spent near the window, frequency of window approaches, duration of window visits, rate of walking near the window, line crossings outside the window area, pecking the window, and crowing were measured. Subjects responded similarly in the presence of male and female stimulus birds during the first test block but discriminated the sex of conspecifics after noncopulatory exposures to males. The discrimination was evident in less locomotion and less crowing in the presence of females as compared with males and, as predicted by a discrimination learning hypothesis, developed largely through a change in responding to males.     

Diffusion of novel foraging behaviour in Amazon parrots through social learning

While social learning has been demonstrated in species across many taxa, the role it plays in everyday foraging decisions is not well understood. Investigating social learning during foraging could shed light on the emergence of cultural variation in different groups. We used an open diffusion experiment to examine the spread of a novel foraging technique in captive Amazon parrots. Three groups were tested using a two-action foraging box, including experimental groups exposed to demonstrators using different techniques and control birds. We also examined the influence of agonistic and pilfering behaviour on task acquisition. We found evidence of social learning: more experimental birds than control birds interacted with and opened the box. The birds were, however, no more likely to use the demonstrated technique than the non-demonstrated one, making local or stimulus enhancement the most likely mechanism. Exhibiting aggression was positively correlated with box opening, whilst receiving aggression did not reduce motivation to engage with the box, indicating that willingness to defend access to the box was important in task acquisition. Pilfering food and success in opening the box were also positively correlated; however, having food pilfered did not affect victims’ motivation to interact with the box. In a group context, pilfering may promote learning of new foraging opportunities. Although previous studies have demonstrated that psittacines are capable of imitation, in this naturalistic set-up there was no evidence that parrots copied the demonstrated opening technique. Foraging behaviour in wild populations of Amazons could therefore be facilitated by low-fidelity social learning mechanisms.     

Life history and social learning: megapode chicks fail to acquire feeding preferences from conspecifics

Australian brush-turkeys, Alectura lathami, are birds with an unusual life history: The young receive no parental care and first encounter conspecifics at an unpredictable age. Brush-turkey chicks that were 3-4 days old were presented with a robot model that appeared to feed from a distinctively colored dish. In control training trials, chicks saw a robot standing next to a different dish and scanning from side to side. Chicks expressed a strong tendency to feed from dishes of the type indicated by the pecking robot, but this effect proved ephemeral. Brush-turkeys hence appear to show no social learning under conditions that inculcate stable preferences in other galliforms such as chickens, suggesting that life history plays an important role in the evolution of learning.     

Time-space learning in homing pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>): orientation to an artificial light source

Time-space learning reflects an ability to represent in memory event-stimulus properties together with the place and time of the event; a capacity well developed in birds. Homing pigeons were trained in an indoor octagonal arena to locate one food goal in the morning and a different food goal in the late afternoon. The goals differed with respect to their angular/directional relationship to an artificial light source located outside the arena. Further, the angular difference in reward position approximated the displacement of the sun's azimuth that would occur during the same time period. The experimental birds quickly learned the task, demonstrating the apparent ease with which birds can adopt an artificial light source to discriminate among alternative spatial responses at different times of the day. However, a novel midday probe session following successful learning revealed that the light source was interpreted as a stable landmark and not as a surrogate sun that would support compass orientation. Probe sessions following a phase shift of the light–dark cycle revealed that the mechanism employed to make the temporal discrimination was prevailingly based on an endogenous circadian rhythm and not an interval timing mechanism.     

NR2B downregulation in a forebrain region required for avian vocal learning is not sufficient to close the sensitive period for song learning

The neural changes that limit the sensitive period for avian song development are unknown, but neurons in a forebrain region critical for song learning, the lMAN, exhibit experience-driven changes in NMDAR subunit expression that could regulate sensitive period closure. Specifically, NR2B levels in lMAN decrease during song acquisition, potentially reducing synaptic plasticity by decreasing NMDAR EPSC duration and/or affecting NMDAR-coupled intracellular cascades. While rearing birds in isolation extends the sensitive period and also delays the developmental changes in NR2B expression and NMDAR physiology, recent work indicates that a transition to faster NMDAR currents does not preclude further song learning. However, NR2B mRNA expression in isolates remains elevated beyond the age at which NMDAR currents shorten, leaving open the possibility that NR2B levels regulate closure of the sensitive period through effects other than those mediated by NMDAR current duration. To determine whether the experience-driven decrease in NR2B expression in lMAN closes the sensitive period, we promoted this change in gene expression either by treating isolation-reared zebra finches briefly with testosterone (T-isolates) or by allowing males limited access to conspecific song (pre-exposed isolates). We then assessed if these birds could acquire song from tutors after the normal close of the sensitive period. Despite a normal decline in NR2B expression, T-isolate and pre-exposed isolate birds learned tutor songs heard from d65-90, while normally reared birds did not. These findings suggest that the normal decline in NR2B expression with lMAN is not sufficient for sensitive period closure.     

Social learning by following: an analysis

Learning by “following”, probably a common means by which behaviors are socially transmitted from adults to young in many species, was analyzed. Pigeons first learned to eat from a human hand. When the hand then approached an operant key and pecked it, the pigeons followed and quickly learned to do the same, thereby demonstrating social learning. When the hand only led the birds to the area of the key, without demonstrating the key-peck response, the birds learned as rapidly as with a key-peck demonstration. Birds also learned, but less reliably and more slowly, when they could observe the hand's responses but were constrained and unable to follow. “Following” was also shown to engender very rapid learning of a more complex, two-member response chain.     

Sociality and social learning in two species of corvids: the pinyon jay (Gymnorhinus cyanocephalus) and the Clark's nutcracker (Nucifraga columbiana)

The hypothesis that social learning is an adaptive specialization for social living predicts that social species should learn better socially than they do individually, but that nonsocial species should not exhibit a similar enhancement of performance under social learning conditions. The authors compared individual and social learning abilities in 2 corvid species: the highly social pinyon jay (Gymnorhinus cyanocephalus) and the less social Clark's nutcracker (Nucifraga columbiana). The birds were tested on 2 different tasks under individual and social learning conditions. Half learned a motor task individually and a discrimination task socially; the other half learned the motor task socially and the discrimination task individually. Pinyon jays learned faster socially than they did individually, but nutcrackers performed equally well under both learning conditions. Results support the hypothesis that social learning is an adaptive specialization for social living in pinyon jays.     

Transfer of matching-to-sample in pigeons

In Experiment I, pigeons were first trained on simultaneous matching-to-sample with either color stimuli or form stimuli, and then shifted to stimuli on the other dimension. Matching performance in the first session with stimuli on a given dimension was not affected by prior matching training with stimuli on the other dimension. However, in the first six color-matching sessions pooled, birds with prior form-matching training performed significantly better than birds without any prior matching training. In Experiment II, birds with experience matching both colors and forms in separate sessions were tested with novel stimulus configurations involving either novel stimuli or novel combinations of familiar colors and forms. Matching performance was not affected by novel stimulus configurations, except that performance dropped to a chance level or below when the standard stimulus was novel. In Experiments II, III, and IV, three of four tests did not show any effect of prior reinforcement of pecks at a novel stimulus, presented alone, on subsequent matching of that stimulus. The results were interpreted as indicating that matching performance in pigeons depends on the learning of stimulus-response chains involving the specific stimuli employed during training. An incidental observation in Experiments I and II was that there were typically more excess pecks at the standard stimulus during form-matching sessions than during color-matching sessions, which may be related to the fact that form matching is more difficult than color matching.     

Self-regulated learning of a natural category: Do people interleave or block exemplars during study?

Despite decades of research focused on the representation of concepts, little is known about the influence of self-regulatory processes when learning natural categories. Such work is vital, as many contexts require self-regulation when we form complex concepts. Previous research has demonstrated that interleaving, as compared to blocking, can improve classification. Thus, as an initial step to explore self-regulated learning of natural concepts, we evaluated whether people chose to block or interleave their practice. According to the search-for-differences hypothesis, people attempt to identify features of birds that distinguish one category (i.e., bird family) from another, and hence should interleave their study. According to the search-for-similarities hypothesis, people attempt to identify features that indicate inclusion into a single category, and hence are expected to block their study. To evaluate these hypotheses, we had participants learn exemplar birds (e.g., Song Sparrow) with their respective bird families (e.g., Sparrow) by selecting the order in which to study bird families. Across four experiments, different formats for selecting exemplars for study were used, so as to provide converging evidence for how participants regulated their learning. Participants overwhelmingly preferred to block their study, even though interleaving is normatively better for learning.     

Factors influencing song development in the white-crowned sparrow (Zonotrichia leucophrys)

Nestling white-crowned sparrows (Zonotrichia leucophrys nuttalli) were hand-reared in sound-isolation chambers under a variety of conditions. The songs of total isolates were compared with songs of birds tutored with song, and the number of inputs sufficient for a bird to produce a normal song was explored. The flexibility of the song learning system was investigated with a range of tape-recorded tutor songs: alien dialects, alien subspecies, alien species, alternating alien dialects, and an aberrant song. Adult songs were obtained for 40 males and 7 testosterone-injected females. All of the tutor songs could be learned. Also, some birds learned elements of an alien species' song. Birds tutored with two songs copied one or the other, were bilingual, or sang a hybrid of the two. No bird presented with fewer than 120 songs learned the tutor song; 2 birds tutored with 252 songs copied the tutor song. It is concluded that the song learning system is quite flexible, hat the results obtained with tape-tutors are very different from those with social tutors, and that there may be an interaction between total number of song inputs and the number presented on a single day. Some implications of these data for physiological mechanisms and the possible functional significance of the acquisition system are discussed.