Summation in the chimpanzee (Pan troglodytes).

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.