Groundbreaking were the experiments carried out by German physician Franz Külbs (1875-1964) between 1904 and 1906 in Kiel. He regularly submitted dogs from the same litter to treadmill training of varying intensity. During the dissection of the animals afterwarts he found an enlargement of the heart, the kidneys, the liver and other organs.
The first scientific studies also took place in water polo. This photo from The New York Tribune tells us that American physician Philip Bovier Hawk (1874-1966) conducted experiments on the water polo players at the University of Pennsylvania, noting that:
"a three minute water polo game produces a larger number of red blood cells than any other sport ...".
Bernarr Macfadden (1868-1955) was an influential American advocate of body culture, a combination of bodybuilding with nutritional and health theories. He also founded the publisher Macfadden Publications. He recommended a minimalist lifestyle based on the time spent in nature, daily heavy physical exertion, abstinence from alcohol, tea, coffee and white bread.
Macfadden sold a wall-mounted muscle developer developed by him and founded Physical Culture in 1899, one of the first muscular magazines. In 1903 he organized the first American bodybuilding competition.
In 1921 and 1922, similar contests marked the triumph for Charles Atlas (1893-1972), the American icon of body culture. In 1935 the publication empire of Macfadden had 35 million readers. In 1955 he died as a multimillionaire.
The book 'Health, Strength & Power' by the American physician Dudley Sargent (1849-1924) was one of the first books on 'physical fitness'.
Danish physiologists August Krogh (1874-1949) and Christian Bohr (1855-1911) published work demonstrating that when carbon dioxide binds to haemoglobin, it decreases the affinity of haemoglobin for oxygen, thus promoting unloading of O2, ‘the Bohr Effect’. Two years later they showed that oxygen passes from alveolus to capillary via passive diffusion.