Response strategies during acquisition of serial learning in pigeons (Columba livia)

Pigeons were exposed on fixed duration sessions to four chromatic stimuli and trained to perform a 2-item sequence. A detailed analysis of incorrect responses performed on each trial prior to completion of the sequence led to distinguish three learning stages. Already on initial sessions animals responded significantly more frequently to relevant stimuli than to irrelevant ones indicating stimulus control by salience. During an intermediary stage, responding to item 2 increased significantly and animals started an increasing number of trials with item 2, suggesting stimulus control by recency. On the final session the correct sequence outnumbered significantly the reverse sequence, suggesting control by order, while recency and salience effects were present to a lesser extent. Repeated pecks, i.e. consecutive pecks on the same cue, occurred on the four cues on the first session and decreased significantly on the last session for irrelevant colours. In addition, when key pecks were followed by a brief sound signal, repeats on relevant stimuli decreased while efficient colour pecks (i.e. when repeats were ignored) on these stimuli increased. These findings are discussed in the light of those obtained in traditional chaining paradigms.     

Attention to uninformative features expressed by pigeons in a duration matching-to-sample task

Pigeons were trained on a matching-to-sample task in which they had to respond to a different choice stimulus following the same durations (2 or 10 s) of two different signals. The duration signals consisted of a white light presented from the ceiling and a red light presented from the front wall. Subsequent test performance indicated that matching accuracy declined (1) when the set of choice stimuli following a duration signal differed from the set presented during training, and (2) when the color or location of the duration signal was changed from values used during training. These results are discussed in terms of attention to uninformative features of a visual stimulus.     

Matching-to-sample by pigeons: the dissociation of comparison choice frequency from the probability of reinforcement

It has been proposed that comparison choice in matching-to-sample should depend on two factors, the relative probability of reinforcement associated with each of the comparison stimuli and the conditional probability of each comparison stimulus being correct given presentation of one of the samples. DiGian and Zentall [DiGian, K.A., Zentall, T.R., 2007. Matching-to-sample in pigeons: in the absence of sample memory, sample frequency is a better predictor of comparison choice than the probability of reinforcement for comparison choice. Learn Behav. 35, 242-261] have shown that sample frequency together with the probability of choosing each of the comparison stimuli in training can influence comparison choice when delays are introduced, when the number of reinforcements associated with each of the comparison stimuli is equated. Furthermore, Zentall and Clement [Zentall, T.R., Clement, T.S., 2002. Memory mechanisms in pigeons: Evidence of base-rate neglect. J. Exp. Psych.: Anim. Behav. Proc. 28, 111-115] have found that sample frequency can affect comparison choice when delays are introduced independently of the number of choices of each of the comparison stimuli in training and the number of reinforcements associated with each of the comparison stimuli is equated. In the present experiment we found that the probability of choosing each of the comparison stimuli in training can affect comparison choice when delays are introduced, independently of sample frequency and when the number of reinforcements associated with each of the comparison stimuli is equated. Together, these experiments suggest that when the sample is not available, there is a partial dissociation between comparison choice and the probability of reinforcement associated with each of the comparison stimuli.     

Learning multiple routes in homing pigeons

The aerial lifestyle of central-place foraging birds allows wide-ranging movements, raising fundamental questions about their remarkable navigation and memory systems. For example, we know that pigeons (Columba livia), long-standing models for avian navigation, rely on individually distinct routes when homing from familiar sites. But it remains unknown how they cope with the task of learning several routes in parallel. Here, we examined how learning multiple routes influences homing in pigeons. We subjected groups of pigeons to different training protocols, defined by the sequence in which they were repeatedly released from three different sites, either sequentially, in rotation or randomly. We observed that pigeons from all groups successfully developed and applied memories of the different release sites (RSs), irrespective of the training protocol, and that learning several routes in parallel did not impair their capacity to quickly improve their homing efficiency over multiple releases. Our data also indicated that they coped with increasing RS uncertainty by adjusting both their initial behaviour upon release and subsequent homing efficiency. The results of our study broaden our understanding of avian route following and open new possibilities for studying learning and memory in free-flying animals.     

Prototype identification and categorization of incomplete figures by pigeons

Previous research has shown that pigeons, unlike humans, cannot identify prototypes after training with corrupted exemplars. In the present experiment, pigeons were initially trained to discriminate between examples of a degraded square and a degraded triangle. The pigeons then received a transfer test in extinction comprised of novel examples of the degraded square and triangle, and the prototype square and triangle. The pigeons successfully categorized both the novel examples of the degraded figures and the underlying prototypes. These results suggest that aspects of categorization and prototype identification may be shared across species.     

Effects of prefeeding, extinction, and distraction during sample and comparison presentation on sensitivity to reinforcer frequency in matching to sample

The present experiment examined the effects of several test manipulations on discrimination, accuracy and sensitivity to reinforcer frequency in a conditional discrimination. Four pigeons responded on a multiple schedule of matching to sample procedures in which the reinforcer-frequency ratio for correct comparison choice responding was varied across components within session from 1:9 to 9:1. Following stability, the effects of prefeeding, extinction, and distraction during sample and comparison presentation were assessed. Discrimination accuracy decreased under prefeeding, extinction, and distraction during sample presentation. Sensitivity to reinforcer frequency decreased under prefeeding and extinction. Decreases in sensitivity were positively related to decreases in discrimination accuracy. The decreases in discrimination accuracy and sensitivity under prefeeding and extinction are interpreted as being due to decreases in attending to the sample and comparison stimuli, respectively, possibly mediated by motivational effects of these manipulations. This interpretation is consistent with current conceptualizations of the contingencies that govern conditional-discrimination performance.     

Dual synaptic plasticity in the hippocampus: Hebbian and spatiotemporal learning dynamics

We assume that Hebbian learning dynamics (HLD) and spatiotemporal learning dynamics (SLD) are involved in the mechanism of synaptic plasticity in the hippocampal neurons. While HLD is driven by pre- and postsynaptic spike timings through the backpropagating action potential, SLD is evoked by presynaptic spike timings alone. Since the backpropagation attenuates as it nears the distal dendrites, we assume an extreme case as a neuron model where HLD exists only at proximal dendrites and SLD exists only at the distal dendrites. We examined how the synaptic weights change in response to three types of synaptic inputs in computer simulations. First, in response to a Poisson train having a constant mean frequency, the synaptic weights in HLD and SLD are qualitatively similar. Second, SLD responds more rapidly than HLD to synchronous input patterns, while each responds to them. Third, HLD responds more rapidly to more frequent inputs, while SLD shows fluctuating synaptic weights. These results suggest an encoding hypothesis in that a transient synchronous structure in spatiotemporal input patterns will be encoded into distal dendrites through SLD and that persistent synchrony or firing rate information will be encoded into proximal dendrites through HLD.     

Intermodal transfer from a visual to an auditory discrimination using an errorless learning procedure

Errorless learning is a technique developed by Terrace [Terrace, H.S., 1963a. Discrimination training with and without “errors”. J. Exp. Anal. Behav. 6, 1-27] to train stimulus discriminations with few or no errors. In the first replication of the original findings, errorless learning was also shown to transfer successfully between two visual discriminations without errors [Terrace, H.S., 1963b. Errorless transfer of a discrimination across two continua. J. Exp. Anal. Behav. 6, 223-232]. In the present experiment, we extended the errorless learning procedure to an intermodal transfer, from a discrimination between red and green colors to a discrimination between high and low tones. The pigeons were divided into two groups: an Experimental Group, which learned both discriminations through errorless learning, and a Control Group, which learned them through trial-and-error. Results showed that pigeons from the Experimental Group learned the red-green discrimination with significantly fewer errors than the Control Group and that errorless learning is effective in transferring from a visual to an auditory discrimination.     

刺激家鸽中脑丘间复合体背内侧核诱发叫声与其自鸣声的比较

利用计算机声谱分析技术比较了家鸽刺激中脑丘间复合体背内侧核(DM)诱发叫声和其正常自鸣声。家鸽的单次自鸣声“di·Gu—”,包括前奏、高潮声和尾声,其主频率和相对幅值都呈明显的逐步递增、平稳和逐步下降过程。高潮声平稳期的载波主频率(318Hz)所表征的主音调比前奏和尾声的主频率(239Hz)分别平均高4.9个半音,相对幅值分别平均高24.4dB和13.2dB,品质因数(Q6dB)分别增高1.8倍和2.8倍。随着刺激电压的增大和减小,家鸽单次鸣声持续时间呈显著的线性递减和递增。诱发叫声的主频率显著性低于自鸣声,声图中有1-2个陪音。本实验为阐明非鸣禽发声调控提供声学特征上的依据     

Evaporative and excretory water loss during free flight in pigeons

Body water conservation is important in flying birds because the very high metabolic demands and heat dissipation requirements during flight depend on plasma-volume integrity. Wind tunnel experiments and theoretical model predictions show that evaporative water loss (EWL) depends on air temperature (T _a) and water vapor density (ρ_a), but these relationships have not been examined in free-flying birds. The contribution of excretory water loss to the total water loss of a flying bird is thought to be negligible but this assumption is untested. To study the dependence of water losses on environmental conditions in free-flying birds and to quantify the contribution of excretory water loss to total water loss, we estimated evaporative and excretory water losses in 16 trained, free-flying tippler pigeons (Columba livia, 250–340 g). We collected excreta by attaching a light latex, water-impermeable receptacle around each bird's vent. By gravimetry, we measured evaporative and excretory water losses of birds for eight flights at different T _as and compared flying to resting (control) birds for two of these flights. EWL was constant with respect to T _a when less than 15 °C, and increased with increasing T _a above 19 °C, indicating that evaporative cooling was invoked when the heat load increased. EWL increased with increasing ρ_a, possibly due to the strong correlation between ρ_a and T _a. Excretory water loss was independent of ρ_a or T _a and averaged almost 10% of the total water loss. Measurements of EWL made on pigeons during wind tunnel experiments and previous free-flight studies are consistent with our free-flight measurements made at similar T _a s. Accepted: 13 April 1999     

Computational Consequences of Temporally Asymmetric Learning Rules: I. Differential Hebbian Learning

Temporally asymetric learning rules governing plastic changes in synaptic efficacy have recently been identified in physiological studies. In these rules, the exact timing of pre- and postsynaptic spikes is critical to the induced change of synaptic efficacy. The temporal learning rules treated in this article are approximately antisymmetric; the synaptic efficacy is enhanced if the postsynaptic spike follows the presynaptic spike by a few milliseconds, but the efficacy is depressed if the postsynaptic spike precedes the presynaptic spike. The learning dynamics of this rule are studied using a stochastic model neuron receiving a set of serially delayed inputs. The average change of synaptic efficacy due to the temporally antisymmetric learning rule is shown to yield differential Hebbian learning. These results are demonstrated with both mathematical analyses and computer simulations, and connections with theories of classical conditioning are discussed.     

Behavioral research in pigeons with ARENA: An Automated Remote Environmental Navigation Apparatus

Three experiments established the effectiveness of an Automated Remote Environmental Navigation Apparatus (ARENA) developed in our lab to study behavioral processes in pigeons. The technology utilizes one or more wireless modules, each capable of presenting colored lights as visual stimuli to signal reward and of detecting subject peck responses. In Experiment 1, subjects were instrumentally shaped to peck at a single ARENA module following an unsuccessful autoshaping procedure. In Experiment 2, pigeons were trained with a simultaneous discrimination procedure during which two modules were illuminated different colors; pecks to one color (S+) were reinforced while pecks to the other color (S−) were not. Pigeons learned to preferentially peck the module displaying the S+. In Experiment 3, two modules were lit the same color concurrently from a set of six colors in a conditional discrimination task. For three of the colors pecks to the module in one location (e.g., upper quadrant) were reinforced while for the remaining colors pecks at the other module (e.g., lower quadrant) were reinforced. After learning this discrimination, the color-reinforced location assignments were reversed. Pigeons successfully acquired the reversal. ARENA is an automated system for open-field studies and a more ecologically valid alternative to the touchscreen.     

Learning to speciate: The biased learning of mate preferences promotes adaptive radiation

Bursts of rapid repeated speciation called adaptive radiations have generated much of Earth's biodiversity and fascinated biologists since Darwin, but we still do not know why some lineages radiate and others do not. Understanding what causes assortative mating to evolve rapidly and repeatedly in the same lineage is key to understanding adaptive radiation. Many species that have undergone adaptive radiations exhibit mate preference learning, where individuals acquire mate preferences by observing the phenotypes of other members of their populations. Mate preference learning can be biased if individuals also learn phenotypes to avoid in mates, and shift their preferences away from these avoided phenotypes. We used individual‐based computational simulations to study whether biased and unbiased mate preference learning promotes ecological speciation and adaptive radiation. We found that ecological speciation can be rapid and repeated when mate preferences are biased, but is inhibited when mate preferences are learned without bias. Our results suggest that biased mate preference learning may play an important role in generating animal biodiversity through adaptive radiation.     

Metabolic responses of homing pigeons to flight and subsequent recovery

This study examines metabolic changes occurring during short to endurance flights and during subsequent recovery in free-flying pigeons, in particular the change towards lipid utilization with increasing flight duration, lipid supply to the flight muscles, protein utilization and the time needed to metabolically recover. Eight plasma metabolite concentrations were measured in homing pigeons released from sites 20–200 km from the loft (0.3–4.8 h flight duration) just after landing and after keeping birds fasting at rest for 30 and 60 min, respectively, after their return. Birds kept in the loft fasting at rest were used as controls. Plasma free fatty acid and glycerol concentrations increased rapidly with flight duration and leveled off after about 1.5 h. This indicates a marked change towards a high and stable lipid utilization from adipose tissues within 1–2 h of flight. Plasma triglyceride levels and very-low-density lipoproteins were decreased after short flights, but subsequently regained or surpassed fasting levels at rest. This indicates that re-esterification of free fatty acids and delivery as very-low-density lipoproteins to the flight muscles to circumvent constraints of fatty acid supply, as described previously for small passerines, is not as significant in the pigeon which has a much lower mass-specific energy rate. An initial increase in plasma glucose levels and a transient decrease to fasting levels at rest was observed and may reflect the initial use and subsequent exhaustion of glycogen stores. Contrary to other birds and mammals, -hydroxy-butyrate levels increased markedly with flight duration. This may suggest a more important sparing of carbohydrates and protein as gluconeogenic precursors in the pigeon than in other species. Plasma uric acid levels increased linearly up to about 4 h flight duration. This indicates an accelerated protein breakdown during flight which may primarily serve to deliver amino acids as glucogenic precursors and citrate cycle intermediates. With increasing flight duration, the energy sources change from an initial phase based primarily on carbohydrates to a lipid-based endurance phase. It is discussed whether this metabolic change depends on the level of power output or the performed work (energy spent) since the start of flight. During the first hour of recovery, most metabolites reached or approached fasting levels at rest, indicating a marked reduction in lipolysis and protein breakdown. -hydroxy-butyrate levels remained at flight levels and glucose levels increased slightly, indicating a restoration of glycogen stores.Abbreviations VLDL very-low-density lipoproteins - FFA free fatty acids     

An experimental comparison of human social learning strategies: payoff-biased social learning is adaptive but underused

Analytical models have identified a set of social learning strategies that are predicted to be adaptive relative to individual (asocial) learning. In the present study, human participants engaged in an ecologically valid artifact-design task with the opportunity to engage in a range of social learning strategies: payoff bias, conformity, averaging and random copying. The artifact (an arrowhead) was composed of multiple continuous and discrete attributes which jointly generated a complex multimodal adaptive landscape that likely reflects actual cultural fitness environments. Participants exhibited a mix of individual learning and payoff-biased social learning, with negligible frequencies of the other social learning strategies. This preference for payoff-biased social learning was evident from the initial trials, suggesting that participants came into the study with an intrinsic preference for this strategy. There was also a small but significant increase in the frequency of payoff-biased social learning over sessions, suggesting that strategy choice may itself be subject to learning. Frequency of payoff-biased social learning predicted both absolute and relative success in the task, especially in a multimodal (rather than unimodal) fitness environment. This effect was driven by a minority of hardcore social learners who copied the best group member on more than half of trials. These hardcore social learners were also above-average individual learners, suggesting a link between individual and social learning ability. The lower-than-expected frequency of social learning may reflect the existence of information producer–scrounger dynamics in human populations.     

Learning outdoors: male lizards show flexible spatial learning under semi-natural conditions

Spatial cognition is predicted to be a fundamental component of fitness in many lizard species, and yet some studies suggest that it is relatively slow and inflexible. However, such claims are based on work conducted using experimental designs or in artificial contexts that may underestimate their cognitive abilities. We used a biologically realistic experimental procedure (using simulated predatory attacks) to study spatial learning and its flexibility in the lizard Eulamprus quoyii in semi-natural outdoor enclosures under similar conditions to those experienced by lizards in the wild. To evaluate the flexibility of spatial learning, we conducted a reversal spatial-learning task in which positive and negative reinforcements of learnt spatial stimuli were switched. Nineteen (32%) male lizards learnt both tasks within 10 days (spatial task mean: 8.16 ± 0.69 (s.e.) and reversal spatial task mean: 10.74 ± 0.98 (s.e.) trials). We demonstrate that E. quoyii are capable of flexible spatial learning and suggest that future studies focus on a range of lizard species which differ in phylogeny and/or ecology, using biologically relevant cognitive tasks, in an effort to bridge the cognitive divide between ecto- and endotherms.     

Superior abstract-concept learning by Clark's nutcrackers (Nucifraga columbiana)

The ability to learn abstract relational concepts is fundamental to higher level cognition. In contrast to item-specific concepts (e.g. pictures containing trees versus pictures containing cars), abstract relational concepts are not bound to particular stimulus features, but instead involve the relationship between stimuli and therefore may be extrapolated to novel stimuli. Previous research investigating the same/different abstract concept has suggested that primates might be specially adapted to extract relations among items and would require fewer exemplars of a rule to learn an abstract concept than non-primate species. We assessed abstract-concept learning in an avian species, Clark''s nutcracker (Nucifraga columbiana), using a small number of exemplars (eight pairs of the same rule, and 56 pairs of the different rule) identical to that previously used to compare rhesus monkeys, capuchin monkeys and pigeons. Nutcrackers as a group (N = 9) showed more novel stimulus transfer than any previous species tested with this small number of exemplars. Two nutcrackers showed full concept learning and four more showed transfer considerably above chance performance, indicating partial concept learning. These results show that the Clark''s nutcracker, a corvid species well known for its amazing feats of spatial memory, learns the same/different abstract concept better than any non-human species (including non-human primates) yet tested on this same task.