The calculating brain

The human brain possesses neural networks and mechanisms enabling the representation of numbers, basic arithmetic operations, and mathematical reasoning. Without the ability to represent numerical quantity and perform calculations, our scientifically and technically advanced culture would not exist. However, the origins of numerical abilities are grounded in an intuitive understanding of quantity deeply rooted in biology. Nevertheless, more advanced symbolic arithmetic skills require a cultural background with formal mathematical education. In the past two decades, cognitive neuroscience has seen significant progress in understanding the workings of the calculating brain through various methods and model systems. This review begins by exploring the mental and neuronal representations of nonsymbolic numerical quantity and then progresses to symbolic representations acquired in childhood. During arithmetic operations (addition, subtraction, multiplication, and division), these representations are processed and transformed according to arithmetic rules and principles, leveraging different mental strategies and types of arithmetic knowledge that can be dissociated in the brain. Although it was once believed that number processing and calculation originated from the language faculty, it is now evident that mathematical and linguistic abilities are primarily processed independently in the brain. Understanding how the healthy brain processes numerical information is crucial for gaining insights into debilitating numerical disorders, including acquired conditions like acalculia and learning-related calculation disorders such as developmental dyscalculia.