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UEXKÜLL, J. Jakob von Uexküll (1864, Keblaste [now Mihkli], Estonia-1944, Capri, Italy) was a biologist, and the founder of biosemiotics. He studied zoology at the University of Tartu (then Dorpat), Estonia, from 1884 to 1889; after that he worked at the Institute of Physiology of the University of Heidelberg in the group led by Wilhelm Kühne (1837-1900), and at the Zoological Station in Naples. In 1907 he was given an honorary doctorate from the University of Heidelberg for his studies in the field of muscular physiology and tonus. One of his results from those years became known as Uexküll’s law, which is probably one of the first formulations of the principle of negative feedback occurring inside an organism. His later work was devoted to the problem of how living beings subjectively perceive their environment, how they build the inner model of the world, and how this model is linked to their behaviour. He introduced the term Umwelt (1909) to denote the subjective world of an organism. This is the notion according to which Uexküll is most frequently cited in the contemporary literature. Uexküll developed a specific method of the experimental study of behaviour which he termed ‘Umwelt-research’. Between 1927 and 1939, Uexküll was the director of the Institut für Umweltforschung (also founded by him) at the University of

Hamburg, spending his summers with his family on Puhtu peninsula (western coast of Estonia) in his summer cottage, where he wrote many of his works (Brock 1934a, 1934b: G. v. Uexküll 1964).