Relative numerousness judgment and summation in young, middle-aged, and older adult orangutans (Pongo pygmaeus abelii and Pongo pygmaeus pygmaeus)

The ability to select the larger of two quantities ranging from 1 to 5 (relative numerousness judgment [RNJ[) and the ability to select the larger of two pairs of quantities with each pair ranging from 1 to 8 (summation) were evaluated in young, middle-aged, and older adult orangutans (7 Pongo pygmaeus abelii and 2 Pongo pygmaeus pygmaeus). Summation accuracy and RNJ were similar to those of previous reports in apes; however, the pattern of age-related differences with regard to these tasks was different from that previously reported in gorillas. Older orangutans were less accurate than the young and middle-aged for RNJ, and summation accuracy was equivalent among age groups. Evidence was found to suggest that the young and middle-aged based their selection of the largest quantity pair on both quantities within each pair during the summation task. These results show a relationship between subject age and the quantitative abilities of adult orangutans.     

Discrete quantity judgments in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus): the effect of presenting whole sets versus item-by-item

The authors examined quantity-based judgments for up to 10 items for simultaneous and sequential whole sets as well as for sequentially dropped items in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), bonobos (Pan paniscus), and orangutans (Pongo pygmaeus). In Experiment 1, subjects had to choose the larger of 2 quantities presented in 2 separate dishes either simultaneously or 1 dish after the other. Representatives of all species were capable of selecting the larger of 2 quantities in both conditions, even when the quantities were large and the numerical distance between them was small. In Experiment 2, subjects had to select between the same food quantities sequentially dropped into 2 opaque cups so that none of the quantities were ever viewed as a whole. The authors found some evidence (albeit weaker) that subjects were able to select the larger quantity of items. Furthermore, the authors found no performance breakdown with the inclusion of certain quantities. Instead, the ratio between quantities was the best performance predictor. The authors conclude that quantity-based judgments rely on an analogical system, not a discrete object file model or perceptual estimation mechanism, such as subitizing.     

Estimating and operating on discrete quantities in orangutans (Pongo pygmaeus)

This study investigated the ability of 3 male orangutans (Pongo pygmaeus; 1 subadult, 2 adults) to estimate, compare, and operate on 2 sets of small quantities (1-6 cereal bits). Experiment 1 investigated the orangutans' ability to choose the larger of 2 quantities when they were presented successively as opposed to simultaneously, thus being perceptually unavailable at the time of choice. Experiment 2 investigated the orangutans' ability to select the larger quantity after the original quantities were augmented or reduced. Orangutans were capable of selecting the larger of 2 quantities in Experiment 1. There was also some evidence from Experiment 2, albeit weaker, that orangutans may mentally combine quantities (but not dissociate) to obtain the larger of 2 quantities. This study suggests that orangutans use a representational mechanism (especially when comparing quantities) to select the larger of 2 sets of items.     

Quantity estimation and comparison in western lowland gorillas (Gorilla gorilla gorilla)

We investigated the quantity judgment abilities of two adult male western lowland gorillas (Gorilla gorilla gorilla) by presenting discrimination tasks on a touch-screen computer. Both gorillas chose the larger quantity of two arrays of dot stimuli. On some trials, the relative number of dots was congruent with the relative total area of the two arrays. On other trials, number of dots was incongruent with area. The gorillas were first tested with static dots, then with dots that moved within the arrays, and finally on a task where they were required to discriminate numerically larger subsets within arrays of moving dots. Both gorillas achieved above-chance performance on both congruent and incongruent trials with all tasks, indicating that they were able to use number as a cue even though ratio of number and area significantly controlled responding, suggesting that number was not the only relevant dimension that the gorillas used. The pattern of performance was similar to that found previously with monkeys and chimpanzees but had not previously been demonstrated in gorillas within a computerized test format, and with these kinds of visual stimuli.     

Summation and numerousness judgments of sequentially presented sets of items by chimpanzees (Pan troglodytes).

Summation and numerousness judgments by 2 chimpanzees (Pan troglodytes) were investigated when 2 quantities of M&Ms were presented sequentially, and the quantities were never viewed in their totality. Each M&M was visible only before placement in 1 of 2 cups. In Experiment 1, sets of 1 to 9 M&Ms were presented. In Experiment 2, the quantities in each cup were presented as 2 smaller sets (e.g., 2 + 2 vs. 4 + 1). In Experiment 3, the quantities were presented as 3 smaller sets (e.g., 2 + 2 + 3 vs. 3 + 4 + 1). In Experiment 4, an M&M was removed from 1 set before the chimpanzees' selection. In Experiments 1 and 2, the chimpanzees selected the larger quantity on significantly more trials than would be predicted by chance. In Experiments 3 and 4, 1 chimpanzee performed at a level significantly better than chance. Therefore, chimpanzees mentally represent quantity and successfully combine and compare nonvisible, sequentially presented sets of items.     

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, Gorilla gorilla) perform on the reversed reward contingency task II: transfer to new quantities, long-term retention, and the impact of quantity ratios

We tested 6 chimpanzees (Pan troglodytes), 3 orangutans (Pongo pygmaeus), 4 bonobos (Pan paniscus), and 2 gorillas (Gorilla gorilla) in the reversed reward contingency task. Individuals were presented with pairs of quantities ranging between 0 and 6 food items. Prior to testing, some experienced apes had solved this task using 2 quantities while others were totally na?ve. Experienced apes transferred their ability to multiple-novel pairs after 6 to 19 months had elapsed since their initial testing. Two out of 6 na?ve apes (1 chimpanzee, 1 bonobo) solved the task--a proportion comparable to that of a previous study using 2 pairs of quantities. Their acquisition speed was also comparable to the successful subjects from that study. The ratio between quantities explained a large portion of the variance but affected na?ve and experienced individuals differently. For smaller ratios, na?ve individuals were well below 50% correct and experienced ones were well above 50%, yet both groups tended to converge toward 50% for larger ratios. Thus, some apes require no procedural modifications to overcome their strong bias for selecting the larger of 2 quantities.     

Relative quantity judgment by Asian elephants (Elephas maximus)

This study investigated whether Asian elephants can make relative quantity judgment (RQJ), a dichotomous judgment of unequal quantities ordered in magnitude. In Experiment 1, elephants were simultaneously shown two baskets with differing quantities of bait (up to 6 items). In Experiment 2, elephants were sequentially presented with baits, which could not be seen by elephants in their total quantities. The task of elephants was to choose the larger quantity in both experiments. Results showed that the elephants chose the larger quantity with significantly greater frequency. Interestingly, the elephants did not exhibit disparity or magnitude effects, in which performance declines with a smaller difference between quantities in a two-choice task, or the total quantity increases, respectively. These findings appear to be inconsistent with the previous reports of RQJ in other animals, suggesting that elephants may be using a different mechanism to compare and represent quantities than previously suggested for other species.     

Relative quantity judgments in South American sea lions (<Emphasis Type="Italic">Otaria flavescens</Emphasis>)

There is accumulating evidence that a variety of species possess quantitative abilities although their cognitive substrate is still unclear. This study is the first to investigate whether sea lions (Otaria flavescens), in the absence of training, are able to assess and select the larger of two sets of quantities. In Experiment 1, the two sets of quantities were presented simultaneously as whole sets, that is, the subjects could compare them directly. In Experiment 2, the two sets of quantities were presented item-by-item, and the totality of items was never visually available at the time of choice. For each type of presentation, we analysed the effect of the ratio between quantities, the difference between quantities and the total number of items presented. The results showed that (1) sea lions can make relative quantity judgments successfully and (2) there is a predominant influence of the ratio between quantities on the subjects’ performance. The latter supports the idea that an analogue representational mechanism is responsible for sea lions’ relative quantities judgments. These findings are consistent with previous reports of relative quantities judgments in other species such as monkeys and apes and suggest that sea lions might share a similar mechanism to compare and represent quantities.     

A mental number line in human newborns

Humans represent numbers on a mental number line with smaller numbers on the left and larger numbers on the right side. A left‐to‐right oriented spatial–numerical association, (SNA), has been demonstrated in animals and infants. However, the possibility that SNA is learnt by early exposure to caregivers’ directional biases is still open. We conducted two experiments: in Experiment 1, we tested whether SNA is present at birth and in Experiment 2, we studied whether it depends on the relative rather than the absolute magnitude of numerousness. Fifty‐five‐hour‐old newborns, once habituated to a number (12), spontaneously associated a smaller number (4) with the left and a larger number (36) with the right side (Experiment 1). SNA in neonates is not absolute but relative. The same number (12) was associated with the left side rather than the right side whenever the previously experienced number was larger (36) rather than smaller (4) (Experiment 2). Control on continuous physical variables showed that the effect is specific of discrete magnitudes. These results constitute strong evidence that in our species SNA originates from pre‐linguistic and biological precursors in the brain.     

Food and token quantity discrimination in capuchin monkeys (<Emphasis Type="Italic">Cebus apella</Emphasis>)

Quantity discrimination is adaptive in a variety of ecological contexts and different taxa discriminate stimuli differing in numerousness, both in the wild and in laboratory settings. Quantity discrimination between object arrays has been suggested to be more demanding than between food arrays but, to our knowledge, the same paradigm has never been used to directly compare them. We investigated to what extent capuchin monkeys’ relative numerousness judgments (RNJs) with food and token are alike. Tokens are inherently non-valuable objects that acquire an associative value upon exchange with the experimenter. Our aims were (1) to assess capuchins’ RNJs with food (Experiment 1) and with tokens (Experiment 2) by presenting all the possible pair-wise choices between one to five items, and (2) to evaluate on which of the two proposed non-verbal mechanisms underlying quantity discrimination (analogue magnitude and object file system) capuchins relied upon. In both conditions capuchins reliably selected the larger amount of items, although their performance was higher with food than with tokens. The influence of the ratio between arrays on performance indicates that capuchins relied on the same system for numerical representation, namely analogue magnitude, regardless of the type of stimuli (food or tokens) and across both the small and large number ranges.     

Hand preference for food processing in wild western lowland gorillas (Gorilla gorilla gorilla)

Western lowland gorillas (Gorilla gorilla gorilla) have not previously been represented in studies of laterality in wild great apes. The discovery of swampy clearings frequented by gorillas in northern Congo has provided the first opportunity to redress this imbalance. Hand preference data are presented from 33 gorillas in seated and standing postures, covering the procurement and processing of 2 to 4 plant species. Levels of hand preference exhibited were low. When data from all postures and plant species were pooled, 33% of gorillas showed hand preferences in excess of chance. In the standing posture, more gorillas exhibited significant left-hand preferences than right, but an overall population-level bias was not evident.     

Mother-infant interactions in western Lowland Gorillas (Gorilla gorilla gorilla): spatial relationships,communication, and opportunities for social learning

This study investigated mother-infant interactions in lowland gorillas (Gorilla gorilla gorilla) with particular focus on the relative role of mothers and infants in creating situations that are potentially conducive to infant social learning. Eleven gorilla mother-infant dyads were focally observed in weekly 1-hr sessions for 12 months. Spatial relationships were affected by age as well as by ambient temperature. Although the youngest infant was encouraged by its mother to walk and climb, mothers showed little or no encouragement in other contexts. In contrast, infants were quite interested in their mothers' activities, on some occasions repeated their mother's behavior, and actively encouraged their mothers to share food, play, or follow them. These findings suggest that gorilla infants are more active than their mothers in creating situations that are potentially conducive to the acquisition of knowledge or skills.     

Conditioned inhibition produced by extinction-mediated recovery from the relative stimulus validity effect: a test of acquisition and performance models of empirical retrospective revaluation

Empirical retrospective revaluation is a phenomenon of Pavlovian conditioning and human causal judgment in which posttraining changes in the conditioned response (Pavlovian task) or causal rating (causal judgment task) of a cue occurs in the absence of further training with that cue. Two experiments tested the contrasting predictions made by 2 families of models concerning retrospective revaluation effects. In a conditioned lick-suppression task, rats were given relative stimulus validity training, consisting of reinforcing a compound of conditioned stimuli (CSs) A and X and nonreinforcement of a compound of CSs B and X, which resulted in low conditioned responding to CS X. Massive posttraining extinction of CS A not only enhanced excitatory responding to CS X, but caused CS B to pass both summation (Experiment 1) and retardation (Experiment 2) tests for conditioned inhibition. The inhibitory status of CS B is predicted by the performance-focused extended comparator hypothesis (J. C. Denniston, H. I. Savastano, & R. R. Miller, 2001), but not by acquisition-focused models of empirical retrospective revaluation (e.g., A. Dickinson & J. Burke, 1996; L. J. Van Hamme & E. A. Wasserman, 1994).     

Recency judgments as a function of word frequency: A framing effect and frequency misattributions

The effects of word frequency on judgments of recency of item presentation were examined in two experiments. Subjects in Experiment 1 were presented two mixed lists of high- and low-frequency words followed by a list assignment task for recognized items. It was found that subjects were biased toward assigning low-frequency words to the more recently presented list. Subjects in Experiment 2 were presented a single mixed list of high- and low-frequency words followed by either a relative recency of presentation judgment task or a relative primacy of presentation judgment task. Each word pair on the tests contained one high-frequency word and one low-frequency word. It was found that, for the recency judgment task, subjects were biased to select the low-frequency item as having been presented more recently. However, on the parallel primacy judgment task, there were no effects of word frequency; moreover, overall accuracy levels were higher with primacy than with recency instructions. We interpret the effects of word frequency on recency judgments in Experiments 1 and 2 in terms of a misattribution of frequency-related differences in recollection-based recognition. The finding that recency and primacy instructions produced different patterns of results provides further evidence (Flexser & Bower, 1974) for an effect on performance of the way in which the temporal judgment task was framed.     

Recognition is used as one cue among others in judgment and decision making

Three experiments with paired comparisons were conducted to test the non-compensatory character of the recognition heuristic (D. G. Goldstein & G. Gigerenzer, 2002) in judgment and decision making. Recognition and knowledge about the recognized alternative were manipulated. In Experiment 1, participants were presented pairs of animal names where the task was to select the animal with the larger population. In Experiment 2, participants chose the safer 1 out of 2 airlines, and 3 knowledge cues were varied simultaneously. Recognition effects were partly compensated by task-relevant knowledge. The compensatory effects were additive. Decisions were slower when recognition and knowledge were incongruent. In Experiment 3, compensatory effects of knowledge and recognition were found for the city-size task which had originally been used to demonstrate the non-compensatory character of the recognition heuristic. These results suggest that recognition information is not used in an all-or-none fashion but is integrated with other types of knowledge in judgment and decision making.     

Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task

Two important elements in problem solving are the abilities to encode relevant task features and to combine multiple actions to achieve the goal. The authors investigated these 2 elements in a task in which gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) had to use a tool to retrieve an out-of-reach reward. Subjects were able to select tools of an appropriate length to reach the reward even when the position of the reward and tools were not simultaneously visible. When presented with tools that were too short to retrieve the reward, subjects were more likely to refuse to use them than when tools were the appropriate length. Subjects were proficient at using tools in sequence to retrieve the reward.     

Relative validity of contextual and discrete cues

Contextual conditioning during relative validity training was explored in 3 experiments that used an appetitive Pavlovian conditioning preparation with rats. Magazine entries were the conditioned response. In Experiment 1, true-discrimination (TD: AX+, BX-) training generated weaker conditioning of X than did pseudodiscrimination (PD: AX+/-, BX+/-) training. The context showed a similar relative validity effect. Also, both PD training and simple partial reinforcement (X+/-) reduced contextual conditioning more than did unsignaled food, a demonstration of relative validity using partial reinforcement. Experiments 2 and 3 used within-subject and between-subjects designs, respectively, and showed that relative validity was determined by the summation of differences in conditioning to both the common element (X) and the context. Our results are consistent with an attentional model or with a computational comparator model but not with the Rescorla-Wagner (R. A. Rescorla & A. R. Wagner, 1972) model.     

More or less: spontaneous quantity discrimination in the domestic cat

We examined spontaneous quantity discrimination in untrained domestic cats in three food choice experiments. In Experiment 1, we presented the cats with two different quantities of food in eight numerical combinations. Overall, the subjects chose the larger quantity more often than the smaller one, and significantly so when the ratio between the quantities was less than 0.5. In Experiment 2, we presented the cats with two pieces of food in four different size combinations. Again, subjects chose the larger piece above chance, although not in the combination where the largest item was presented. In Experiment 3, a subset of the cats was presented multiple times with two different quantities of food, which were hidden from view. In this case, the cats did not choose the larger quantity more often than the smaller one, suggesting that in the present experiments they mainly used visual cues when comparing quantities. We conclude that domestic cats are capable of spontaneously discriminating quantities when faced with different numbers or sizes of food items, and we suggest why they may not always be motivated to choose the larger quantity. In doing so, we highlight the advantages of testing spontaneous choice behavior, which is more likely to reflect animals’ everyday manner of responding than is the case when training them in order to test their absolute limits of performance which may not always coincide with their daily needs.     

Discrimination of small quantities by fish (redtail splitfin, Xenotoca eiseni)

Discrimination of quantity has been argued to rely on two non-verbal representational systems: an object file system (OFS) for representing small values (≤3–4) and an analog magnitude system (AMS) for representing large magnitudes (>4). Infants’ ability to discriminate 1 versus 2, 1 versus 3, 2 versus 3, but not 1 versus 4 or 2 versus 4 seems to prove the independence of such systems. Here, we show that redtail splitfin fish (Xenotoca eiseni) performed relative quantity estimations preferring to approach the location previously occupied by the larger in number between two groups of conspecifics (no longer visible at test) in sets of 1 versus 2 and 2 versus 3 items, but failed at 3 versus 4 items, thus showing the same set-size limit as infants for discrimination of small quantities. However, when tested with quantities that spanned the boundary of the two systems, that is, 1 versus 4 and 2 versus 4, fish succeeded. These results thus point to either the use of continuous physical variables and/or the use of the AMS also for small numerousness in fish in this task.     

Summation of symbols by pigeons (Columba livia): the importance of number and mass of reward items

In each of 3 experiments, different sets of 4 pigeons (Columba livia) were trained to discriminate between 2 visual symbols that covered wells containing food items that varied in number, mass, or both. In Experiment 1, the symbols were associated with 0, 1, 3, 5, 7, or 9 pieces of grain reward. The pigeons learned to choose the symbol corresponding to the larger reward, and on summation tests, they chose the pair of symbols that summed to the larger total reward. When number of food pellets was varied but mass of reward was held constant in Experiment 2, preference for the larger number symbols failed to appear. When number was held constant and mass was varied in Experiment 3, the pigeons showed a clear preference for the larger mass symbols on single-symbol and summation tests. These findings show that pigeons summate the value of symbols and are more likely to represent symbols by mass of food reward than by number of food items.