Anatomy and physiology

Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject. As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.

As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy. These papers were left to his heir, Francesco Melzi, for publication, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing. The project was left incomplete at the time of Melzi's death more than 50 years later, with only a small amount of the material on anatomy included in Leonardo's Treatise on painting, published in France in 1632. During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.

Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics. He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero. The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.

Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness. Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.

Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.