Abstract: In this research, we asked whether 2 chimpanzee (Pan troglodytes) subjects could reliably sum across pairs of quantities to select the greater total. Subjects were allowed to choose between two trays of chocolates. Each tray contained two food wells. To select the tray containing the greater number of chocolates, it was necessary to sum the contents of the food wells on each tray. In experiments where food wells contained from zero to four chocolates, the chimpanzees chose the greater value of the summed wells on more than 90% of the trials. In the final experiment, the maximum number of chocolates assigned to a food well was increased to five. Choice of the tray containing the greater sum still remained above 90%. In all experiments, subjects reliably chose the greater sum, even though on many trials a food well on the “incorrect” tray held more chocolates than either single well on the “correct” tray. It was concluded that without any known ability to count, these chimpanzees used some process of summation to combine spatially separated quantities. Speculation regarding the basis for summation includes consideration of perceptual fusion of pairs of quantities and subitization.

Abstract: Five chimpanzees with training in counting and numerical skills selected between 2 arrays of different amounts of candy or 2 Arabic numerals. A reversed reinforcement contingency was in effect, in which the selected array was removed and the subject received the nonselected candies (or the number of candies represented by the nonselected Arabic numeral). Animals were unable to maximize reward by selecting the smaller array when candies were used as array elements. When Arabic numerals were substituted for the candy arrays, all animals showed an immediate shift to a more optimal response strategy of selecting the smaller numeral, thereby receiving the larger reward. Results suggest that a response disposition to the high-incentive candy stimuli introduced a powerful interference effect on performance, which was effectively overridden by the use of symbolic representations.

Abstract: Two chimpanzees and a rhesus macaque rapidly learned the ordinal relations between 5 colors of containers (plastic eggs) when all containers of a given color contained a specific number of identical food items. All 3 animals also performed at high levels when comparing sets of containers with sets of visible food items. This indicates that the animals learned the approximate quantity of food items in containers of a given color. However, all animals failed in a summation task, in which a single container was compared with a set of 2 containers of a lesser individual quantity but a greater combined quantity. This difficulty was not overcome by sequential presentation of containers into opaque receptacles, but performance improved if the quantitative difference between sizes was very large.

Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.