LUDWIG, CARL F.W.

born Dec. 29, 1816, Witzenhausen, near Kassel, Hesse-Kassel died April 23, 1895, Leipzig, Ger. in full Carl Friedrich Wilhelm Ludwig a founder of the physicochemical school of physiology in Germany. A professor of physiology at the universities of Marburg (184649), Zrich (184955), Vienna (185565), and Leipzig (186595), Ludwig is best known for his study of the cardiovascular system. He invented (1847) a device known as a kymograph to record changes in arterial blood pressure; a simple stromuhr (1867), or flowmeter, to measure the rate of blood flow through arteries and veins; and a mercurial blood-gas pump for the separation of gases from the blood, which led to an understanding of the role played by oxygen and other gases in the purification of blood. Ludwig was the first to keep animal organs alive in vitro (outside the animal's body) by perfusing frog hearts with a solution approximating the composition of blood plasma (1856); to locate a blood vessel regulatory mechanism in the medulla oblongata (at the base of the brain); and to measure blood pressure in the capillaries. He discovered the depressor and accelerator nerves of the heart and, with the American physiologist Henry Bowditch, formulated (1871) the all-or-none law of cardiac muscle action, stating that the heart muscle, under whatever stimulus, will contract to the fullest extent or not at all. Modern theories of urine and lymph formation stem from Ludwig's paper (1844) on urine secretion, postulating that the surface layer, or epithelium, of the kidney tubules (known as glomeruli) serves as a passive filter in urine production, the rate of which is controlled by blood pressure. He also introduced the measurement of nitrogen in the urine as an indication of the approximate rate of protein metabolism in the entire animal and was first to show that human digestive glands may be influenced by secretory nerves. Ludwig is considered one of the great physiology teachers; nearly 200 of his students, including Bowditch and the American physician William Welch, became prominent scientists.