Parallel versus sequential processing of relational stimulus structures

Organisms are often faced with sets of stimuli bearing specifiable relationships to each other. Experimental data suggest that even animals not suspected of being particularly rational can solve problems involving consistent linear relationships. We examine the information processing required to cope with these and related stimulus structures from a theoretical point of view. We show that both a parallel processing neural network model and a serially processing Turing machine model require minimal complexities to process linear hierarchical structures. When dealing with other relational stimulus structures, the models need differing, greater minimal complexities. Siemann and Delius (1994) report experimental results indicating that both pigeons and humans appear to operate according to the parallel, neural network model we propose here. Further experiments likely to be diagnostic are proposed.     

Functional equivalence in pigeons involving a four-member class

Research suggests that animals are capable of forming functional equivalence relations or stimulus classes of the kind usually demonstrated by humans (e.g., the class defined by an object and the word for that object). In pigeons, such functional equivalences are typically established using many-to-one matching-to-sample in which two samples are associated with one comparison stimulus and two different samples are associated with the other. Evidence for the establishment of functional equivalences between samples associated with the same comparison comes from transfer tests. In Experiment 1, we found that pigeons can form a single class consisting of four members (many-to-one matching) when the alternative class has only one member (one-to-one matching). In Experiment 2, we ruled out the possibility that the pigeons acquired the hybrid one-to-one/many-to-one task by developing a single-code/default coding strategy as earlier research suggested that it might. Thus, pigeons can develop a functional class consisting of as many as four members, with the alternative class consisting of a single member.     

Intradimensional and extradimensional shifts in conditional discrimination

Two groups of pigeons learned a two key conditional discrimination. Color was the conditional stimulus and form the choice stimulus for subjects in one group. Form was the conditional stimulus and color the choice stimulus in the other group. Half the subjects in each group then underwent an intradimensional shift: The conditional stimulus dimension and choice stimulus dimension were unchanged but the correct and incorrect stimulus compounds were reversed. The remaining subjects underwent an extra-dimensional shift in which the conditional stimulus dimension and choice stimulus dimension were reversed. Subjects which experienced an intradimensional shift learned the new conditional discrimination more quickly. It was concluded that subjects followed rules to solve conditional discriminations but also learned the functions of each stimulus dimension.     

Speeding up reaction time with invisible stimuli

Normal subjects react more quickly to a pair of visual stimuli than to a stimulus alone. This phenomenon is known as the redundant signal effect (RSE) and represents an example of divided visual attention in which signal processing is carried out in parallel to the advantage of response speed. A most interesting aspect of this phenomenon is that it can occur when one stimulus in a pair cannot be consciously detected because of hemianopia or unilateral extinction resulting from brain damage. Here, we report that a similar dissociation between visual awareness and visually guided behavior is present in normal subjects who show an RSE even when the luminance of one of a pair of stimuli is below detection threshold. The observed RSE cannot be attributed to probability summation because it violates Miller's race inequality and is likely to be related to neural summation between supra- and subthreshold stimuli. Given that a similar implicit RSE is present in hemispherectomy patients, we hypothesize that the site of this summation might be the superior colliculus (SC).     

Psychological distance to reward in monkeys

A concurrent-chains schedule was employed to examine crab-eating macaques' choice between segmented and unsegmented fixed-interval 30-s schedules. A stimulus change occurred during one of the fixed-interval schedules (segmented schedule), while no stimulus change occurred during the other (unsegmented schedule). The subjects were exposed to three consecutive stimulus-change conditions, in an attempt to explore whether correlation of the stimulus changes with a fixed interval of nonreinforcement affects the preference between the alternatives. When the stimulus change always occurred at a half point of the segmented schedule (the first and third conditions), all subjects preferred the fixed-interval schedule without stimulus changes. This replicated previous findings with pigeons and humans. On the other hand, when the intervals before and after the stimulus change varied on every entry to the segmented schedule with the sum of them being constant (the second condition), the subjects were indifferent to the alternatives. These results are consistent with the view that the preference for unsegmented schedules over segmented ones might be due to aversiveness which the initial component in the segmented schedule possesses as a result of its correlation with a fixed interval of nonreinforcement.     

Mechanisms of same/different concept learning in primates and avians

Mechanisms of same/different concept learning by rhesus monkeys, capuchin monkeys, and pigeons were studied in terms of how these species learned the task (e.g., item-specific learning versus relational learning) and how rapidly they learned the abstract concept, as the training set size was doubled. They had similar displays, training stimuli, test stimuli, and contingencies. The monkey species learned the abstract concept at similar rates and more rapidly than pigeons, thus showing a quantitative difference across species. All species eventually showed full concept learning (novel-stimulus transfer equivalent to baseline: 128-item set size for monkeys; 256-item set for pigeons), thus showing a qualitative similarity across species. Issues of stimulus regularity/symmetry, generalization from item pairs, and familiarity processing were not considered to be major factors in the final performances, converging on the conclusion that these species were increasingly controlled by the sample-test relationship (i.e., relational processing) leading to full abstract-concept learning.     

Olfactory discrimination ability for aliphatic odorants as a function of oxygen moiety

We tested the ability of human subjects to distinguish between aliphatic odorants sharing the same number of carbon atoms but differing in their functional groups. 1-Alcohols, n-aldehydes, 2-ketones and n-carboxylic acids of four, six and eight carbon atoms, respectively, were employed. In a forced-choice triangular test procedure 20 subjects were repeatedly presented with 18 odor pairs and asked to identify the bottle containing the odd stimulus. We found (i) that as a group, the subjects performed significantly above chance level in all tasks and thus were clearly able to discriminate between all odor pairs presented; (ii) marked interindividual differences in discrimination performance, ranging from subjects who were able to significantly distinguish between all 18 odor pairs to subjects who failed to do so with 1/3 of the tasks; (iii) a lack of significant differences in performance between male and female, and between Japanese and German subjects; (iv) that odor pairs that involved 2-ketones and/or n-carboxylic acids were significantly easier to discriminate compared to odor pairs that involved 1-alcohols and/or n-aldehydes, and thus a clear dependence of discriminability on type of functional group; and (v) that aliphatic odorants with eight carbon atoms (irrespective of their oxygen moiety) were significantly more difficult to discriminate from each other compared to substances with four or six carbon atoms. The results suggest that functional groups may be an important determinant of the interaction between stimulus molecule and olfactory receptor in aliphatic substances, and thus may be a molecular property affecting odor quality in a substance class-specific manner.     

Picture – Object Recognition in Kea (Nestor notabilis)

Although pictures are widely used as stimuli in cognitive experiments with both humans and animals, the question of how subjects interpret pictures receives less attention. Gaining a better understanding of this is especially important when working with avian subjects, as their visual anatomy and processing is different from that of humans, and even differs from one avian species to another. Successful testing for picture recognition in birds has been carried out mainly with pigeons, but no such research has been explicitly performed with ‘brainy’ birds like parrots, despite the fact that these have been the subject of exciting cognitive research. This study tested kea (Nestor notabilis) mountain parrots for picture–object recognition using a procedure which required the transfer of a learned discrimination task between pictures and objects. Kea successfully showed both picture‐to‐object and object‐to‐picture transfer and performed at a comparable level when pictures were displayed on a touch screen or as printed photographs.     

Taste qualities of solutions preferred by hamsters

Molecules of diverse chemical structure are sweet to humans and several lines of evidence (genetic, physiological, behavioral) suggest that there may be distinct sweet perceptual qualities. To address how many perceptual categories these molecules elicit in hamsters (Mesocricetus auratus), we studied patterns of generalization of conditioned taste aversions for seven sweeteners: 100 mM sucrose, 320 mM maltose, 32 mM D-phenylalanine, 3.2 mM sodium saccharin, 16 mM calcium cyclamate, 10 mM dulcin and 32 mM sodium m-nitrobenzene sulfonate. Each stimulus was preferred versus water in two-bottle intake tests and stimulated the chorda tympani nerve. For each of seven experimental groups the conditional stimulus (CS) was a sweetener and for the control group the CS was water. Apomorphine.HCl was injected i.p. after a CS was sampled and, after recovery, test stimuli (TS) were presented for 1 h daily. The intake (ml) of each TS consumed by experimental animals was compared with mean TS intake by the control group. Learned aversions for 18/21 stimulus pairs cross-generalized, resulting in a single cluster of generalization patterns for the seven stimuli. Cross-generalization failures (maltose-cyclamate, maltose-sucrose, cyclamate-NaNBS) may be the consequence of particular stimulus features (e.g. salience, cation taste), rather than the absence of a 'sucrose-like' quality. The results are consistent with a single hamster perceptual quality for a diverse set of chemical structures that are sweet to humans.     

Olfactory discrimination ability for aliphatic c6 alcohols as a function of presence, position, and configuration of a double bond

The ability of human subjects to distinguish between aliphatic C6 alcohols differing in presence, position, or configuration (i.e., cis-trans geometry) of a double bond was tested. In a forced-choice triangular test procedure, 20 subjects were repeatedly presented with all 21 binary combinations of the seven stimuli and asked to identify the bottle containing the odd stimulus. I found (a) that as a group, the subjects performed significantly above chance level in all tasks but two and thus were clearly able to discriminate between most of the odor pairs presented; (b) marked interindividual differences in discrimination performance, ranging from subjects who were able to significantly distinguish between all 21 odor pairs to subjects who failed to do so with 10 of the tasks; (c) that odor pairs involving two hexenols were significantly more difficult to discriminate than odor pairs that involved hexanol and one of the hexenols; (d) that odor pairs involving hexenols sharing the same geometry but differing in the position of the double bond by only one carbon atom were significantly more difficult to distinguish than odor pairs that involved hexenols differing by two carbon atoms; (e) that odor pairs involving 4-hexenols were significantly easier to discriminate than 3-hexenols, which, in turn, were significantly easier to distinguish than 2-hexenols; and (f) that odor pairs involving two cis-hexenols were significantly more difficult to discriminate than odor pairs that involved two trans-hexenols. These findings demonstrate that the presence as well as the position and configuration of a double bond affected discriminability in a systematic manner and suggest that these molecular structural features may be important determinants of the interaction between stimulus molecule and olfactory receptor and thus may affect odor quality of aliphatic alcohols.     

Role of stimulus-stimulus pairing in matching-to-sample procedure: cross-species comparison of humans and pigeons

The present experiment examined whether, in a matching-to-sample (MTS) procedure, a relation between two stimuli, a sample and a comparison, could be established as a result of just stimulus–stimulus pairing, even if back up reinforcers were never provided for the conditional relation between the sample and comparison stimuli, but rather only for the comparison stimulus. A procedure called “pseudo matching-to-sample” was used in which, when S1 was presented as a sample stimulus, two comparison stimuli (C1 and C2) were presented, and only responses to C1 were reinforced. Conversely, when S2 was presented, only responses to C3 (and not C4) were reinforced. In other words, organisms experiencing this procedure could discriminate C1 from C2, and C3 from C4, by simple discrimination without regard to the conditional sample stimuli. In order to examine cross-species differences, responding by humans in this procedure was compared to that by pigeons. Although the humans developed a discriminative function for the sample stimuli, that is, the humans’ responding was affected by both the sample stimuli and the reinforcers, responding by the pigeons was affected solely by the reinforcers. These data suggest that, in this procedure, humans (but not pigeons) are able to learn relations among stimuli simply as a result of stimulus–stimulus pairing.     

A slowing effect on visual search by advance information in pigeons (Columba livia): a comparison with humans (Homo sapiens)

A comparative study was conducted to investigate whether the search for a target letter was facilitated when the target and prime (preceding stimulus) letters were identical. Pigeons (Section 2) and human participants (Section 3) were first trained to search for “A” among “Y”s and “E” among “D”s in a condition in which a square shape appeared as the prime (Neutral condition). In subsequent testing, a prime was identical either to the corresponding target (Target-priming condition) or to the distractor (Distractor-priming condition). Humans and pigeons responded differently to the two priming conditions. On early trials, the Target prime facilitated search in humans, reducing reaction times (RTs) to targets. In pigeons, however, RTs were longer with Target primes, suggesting that pre-exposure to target letters may directly inhibit the search for targets in subsequent search displays. Furthermore, pre-exposure to the distractor letters may inhibit the processing of the distractor. On later trials, RTs of humans were faster in both priming conditions than in the Neutral condition, suggesting that expectation of a target facilitated search (“Y” predicted “A” and “D” predicted “E”). In contrast, the pigeons showed no evidence of expectation-based facilitation, with constant slowing effects of the Target prime extending across sessions. Possible mechanisms underlying such a slowing priming effect in pigeons were discussed.     

Modeling convergent ON and OFF pathways in the early visual system

For understanding the computation and function of single neurons in sensory systems, one needs to investigate how sensory stimuli are related to a neuron’s response and which biological mechanisms underlie this relationship. Mathematical models of the stimulus–response relationship have proved very useful in approaching these issues in a systematic, quantitative way. A starting point for many such analyses has been provided by phenomenological “linear–nonlinear” (LN) models, which comprise a linear filter followed by a static nonlinear transformation. The linear filter is often associated with the neuron’s receptive field. However, the structure of the receptive field is generally a result of inputs from many presynaptic neurons, which may form parallel signal processing pathways. In the retina, for example, certain ganglion cells receive excitatory inputs from ON-type as well as OFF-type bipolar cells. Recent experiments have shown that the convergence of these pathways leads to intriguing response characteristics that cannot be captured by a single linear filter. One approach to adjust the LN model to the biological circuit structure is to use multiple parallel filters that capture ON and OFF bipolar inputs. Here, we review these new developments in modeling neuronal responses in the early visual system and provide details about one particular technique for obtaining the required sets of parallel filters from experimental data.     

Relative numerousness judgements in crows and pigeons in the urgent comparison of stimuli earlier linked to different amounts of reinforcement]

The ability of pigeons (Colomba livia, L.) and crows (Corvus corone cornix, L.) was studied to realize urgent numerousness judgement of reinforcement consisting of discrete elements (wheat grains and meal worm larvae, respectively). In the process of preliminary training the birds mastered the information about the conformity of the feeder colour with the definite number (1-9 for pigeons and 5-12 for crows) of reinforcement units at isolated presentation of feeders. In test at presentation of pairs formed from these feeders, pigeons and crows chose the stimulus connected with a greater quantity of reinforcement. In the range of 1-8 units the precision of choice in pigeons depended on absolute and relative differences between comparing values. In crows in the range of 6-12 this dependence was not revealed. The ability to solve the given test is considered as one of manifestations of elementary reasoning.     

Discrimination of cartoons and photographs in pigeons: effects of scrambling of elements

Four groups of pigeons were trained on four visual discrimination tasks using people photographs, pigeon photographs, cartoons of people, and cartoons of pigeons. After completion of the discrimination, the subjects were tested with new stimuli in the same stimulus categories but never used during the training, and scrambled stimuli in which elements of the original stimulus were spatially randomized. The subjects showed generalization to the new stimuli except for the people cartoons. When scrambled stimuli were presented, the subjects showed suppression of responding to the photographs but not to the people cartoon. These results suggest that the pigeons recognized both the elements and spatial arrangement of the photographs but ignored the spatial arrangement of the human cartoons.     

A combined perceptual, physico-chemical, and imaging approach to 'odour-distances' suggests a categorizing function of the Drosophila antennal lobe

How do physico-chemical stimulus features, perception, and physiology relate? Given the multi-layered and parallel architecture of brains, the question specifically is where physiological activity patterns correspond to stimulus features and/or perception. Perceived distances between six odour pairs are defined behaviourally from four independent odour recognition tasks. We find that, in register with the physico-chemical distances of these odours, perceived distances for 3-octanol and n-amylacetate are consistently smallest in all four tasks, while the other five odour pairs are about equally distinct. Optical imaging in the antennal lobe, using a calcium sensor transgenically expressed in only first-order sensory or only second-order olfactory projection neurons, reveals that 3-octanol and n-amylacetate are distinctly represented in sensory neurons, but appear merged in projection neurons. These results may suggest that within-antennal lobe processing funnels sensory signals into behaviourally meaningful categories, in register with the physico-chemical relatedness of the odours.     

Absolute and relational control of a sequential auditory discrimination by pigeons (Columba livia)

Recent evidence indicates that pigeons can readily learn visual discriminations based on both absolute and relational stimulus factors. To examine how these two types of control function in their non-dominant auditory modality, we tested four pigeons in a go/no-go sequential auditory discrimination in which both absolute and relational cues were redundantly available. In this task, sequences of different sounds created from one set of pitches were reinforced, while different sequences created from another set of pitches and any same sequences made from either set of pitches were not. Across three experiments, we independently varied the relative discriminability of the absolute and relational components. The pigeons were consistently and primarily controlled by the absolute fundamental pitch of our notes in all of the experiments, although this was influenced by the range and arrangement of the pitches used in each set. A majority of the pigeons also demonstrated relational control when this component was made more salient. The more robust control exhibited by absolute factors is consistent with the comparative hypothesis that birds in general may have a well-developed aptitude for processing absolute pitch in many auditory settings. The relational control is consistent with our recent evidence of same/different auditory learning by pigeons.     

Preference for a segmented schedule using a brief S+ stimulus correlated with a great delay reduction in humans

Past studies using the concurrent-chain procedure showed that pigeons and humans generally prefer an unsegmented schedule to a segmented schedule. This finding is ostensibly inconsistent with theories of conditioned reinforcement such as delay-reduction theory. In the present study with humans, two changes in the basic segmented schedule were implemented to resolve this inconsistency. The first change was that in the segmented schedule the terminal-link stimulus (S+ stimulus) changed late in the terminal-link, close to reinforcement presentation. The second change was that the presentation of the segmenting stimulus, S+, was brief allowing a reinstatement of the early terminal-link stimulus, which is contiguous with reinforcement. Our data constitute the first demonstration of preference for the segmented schedule when a brief S+ is correlated with a greater reduction in delay to reinforcement.     

Transverse patterning in pigeons

The ability of pigeons to form configural stimulus representations was assessed in two operant discrimination experiments. In Experiment 1 the transverse patterning problem of Spence (1952) was trained. In Phase 1, subjects had to choose stimulus A on A + B — trials; B + C — trials were added in Phase 2. In Phase 3, the first two pairs were combined with C + A — trials. The success of the subjects was simulated by a model assuming that elemental and configural stimulus representations coexist in a stable proportion, even in the phases of the experiment which do not require configural stimulus representations for successful solution. Experiment 2 replicated the first two phases of Experiment 1, but trained A + C — in Phase 3. Comparison of the results of this experiment with simulations of the model showed that elemental and configural stimulus representations coexisted in similar proportions as in Experiment 1, even though they were not necessary for successful task solution.     

Quantitive studies of the reactions to horizontal angular accelerations in axolotls. II. head-turning reflexes in animals with a supernumerary pair of labyrinths.

In axolotls (Ambystoma mexicanun) the labyrinths and the associated parts of the medulla were doubled artificially. In these so-called tandem-heads the vestibular afferent fibres from both labyrinths on one side united within the medulla to form common bundles. The head-turning reflexes following impulse acceleration and during long-lasting acceleration were measured quantitatively and compared with those for normal animals. The form and the time-course of the reactions were almost identical in both groups. Tandem-heads showed a linear relationship between stimulus intensity and reaction strength, parallel to that in normal animals but with a greater reaction for a given stimulus. Consequent to this shift in the relationship, there was a significant decrease in the reaction threshold. The removal of one horizontal semicircular canal in tandem-heads proved that both pairs of labyrinths were functionally connected with the brain. It was suggested that during ontogenesis there exists a kind of specificity in the connexion of vestibular fibres. From the parallel shift of the intensity functions it was concluded that the input from both pairs of labyrinths in tandem-heads is not simply accumulate but compared with a reference parameter, which is also double in tandem-heads.