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1 2 3 4 5 6 7 8 9 Johann Paul Karplus (1866-1936) 10 11 12 13 14 15 16 17 18 19 20 21 Lazaros C. Triarhou 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 ——————————--—— 46 Prof. L.C. Triarhou, MD, PhD 47 Economo-Koskinas Wing for Integrative and Evolutionary Neuroscience 48 49 University of Macedonia 50 Thessaloniki, Greece 51 E-Mail: [email protected] 52 53 54 55 56 57 58 59 60 61 62 63 64 65

Johann Paul Karplus (Fig. 1) was born on 25 October 1866 in the Moravian-Silesian town of 1 2 Troppau (today Opava, Czech Republic). He was the third child, among four boys and two 3 4 5 girls, of Gottlieb Karplus (1836-1887), a Jewish merchant, and Elisabeth Karplus (1841- 6 7 1925). 8 9 10 After graduating from the University of in 1890, Karplus worked as assistant 11 12 physician in internal medicine (1891-1903) under Hermann Nothnagel, psychiatry and 13 14 (1898-1900) under Richard von Krafft-Ebing, neuropathology (1900-1903) under 15 16 17 Heinrich Obersteiner, and physiology (1903-1917) under Siegmund Exner. 18 19 At his alma mater, Karplus was appointed Privatdocent in psychiatry and neurology in 20 21 22 1901, assistant (Titular) professor of neurophysiology and neuropathology in 1909, and 23 24 associate (Extraordinarius) professor in 1914. During World War I, he served as consultant 25 26 27 physician at Rosenhügel Mental Hospital. From 1917 to 1933 he headed the Department of 28 29 Neurology and Psychiatry at Vienna’s General Polyclinic [1]. 30 31 In his first monograph on the human brain [2] Karplus studied the external configuration 32 33 34 of the cerebral hemispheres in 49 brains from 21 families, including fraternal twins, and 35 36 described anatomical similarities in gyral pattern and hemispheric variations among relatives. 37 38 39 The next monograph on cerebral variability and heredity [3] covered the macroscopic pattern 40 41 of the cerebral cortex, brainstem and spinal cord in human, simian, canine, feline, and caprine 42 43 44 brains, also including kindred and twins. An expanded edition [4] placed emphasis on 45 46 development and gender. 47 48 Between 1893 and 1933, Karplus published 80 articles, 50 of them single-authored [6]. 49 50 51 He wrote on progressive paralysis, ophthalmoplegia, hysteria, pseudobulbar paresis, epilepsy, 52 53 cerebellar physiology, Friedreich ataxia, cutaneous sensory disruption in Parkinson disease, 54 55 56 aneurysms of the basal cerebral arteries, migraine, spinal pain mechanisms, familial 57 58 syringomyelia, regulation of perspiration, hyperidrosis in hemiplegia, pyramidal tract 59 60 61 62 63 2 64 65

degeneration after cortical lesions in marmoset and macaque monkeys, and comparative 1 2 anatomy of Australian aborigine and African human brains, noting no differences compared 3 4 5 to the average European. Otto Marburg described the writing style of Karplus as concise, 6 7 clear, matter-of-fact, free of imaginary explanations, and “exemplary for all young 8 9 10 neurologists” [7] . 11 12 Co-authors on the remaining 30 publications included Alexander Spitzer (anatomy of the 13 14 brainstem and experimental lesions of pons and brachium pontis, 1904-1907); Constantin von 15 16 17 Economo (physiology and anatomy of midbrain and lesions of cerebral peduncles in cats and 18 19 macaques, 1909); neurosurgeon Alfred Exner (radiotherapy of malignant brain tumors, 1913); 20 21 22 and Emil Redlich (contused wounds from grenade explosions, traumatic war neuroses, and 23 24 epilepsy as sequel of head injury, 1916-1917). 25 26 27 The milestone discovery credited to Karplus and the Bohemian physiologist Alois Kreidl 28 29 (pupil of Josef Breuer) is the discovery of a region in the posterior hypothalamus which 30 31 contains the mechanism that subserves autonomic responses (‘Karplus-Kreidl center’ of 32 33 34 vegetative function in the older physiological literature). In a prolific collaboration spanning 35 36 over 20 years, the two scientists applied faradic stimulation of the hypothalamus in 37 38 39 anesthetized cats and documented bodily responses reminiscent of those after sympathetic 40 41 nervous discharge. They also described the pathway of the pupillary light reflex. Between 42 43 44 1909 and 1928, the year of Kreidl’s death, they published 8 papers in Pflügers Archiv on the 45 46 ‘Brain and sympathetic’ [8, 9]. In one of these studies they collaborated with Dutch 47 48 cardiologists Simon Hoogerwerf and Willem Einthoven, the latter a Nobel laureate in 1924 49 50 51 for discovering the mechanism of the electrocardiogram. From 1930 until 1933 Karplus 52 53 carried out research on the effects of hypothalamic stimulation on the pituitary together with 54 55 56 Kreidl’s pupil, pharmacologist Oscher Peczenik [6]. 57 58 Based on Karplus and Kreidl’s discoveries, Philip Bard, a pupil of Walter B. Cannon, 59 60 61 62 63 3 64 65

carried out ablation experiments and concluded in 1928 that the posterior hypothalamus 1 2 contains the central locus requisite for “sham rage,” a condition of vigorous sympathoadrenal 3 4 5 activity resembling the behavior of “infuriated” animals [8]. The findings of Karplus and 6 7 Kreidl were confirmed by Stephen W. Ranson of Northwestern University in the 1930s, by 8 9 10 Detlev Wulf Bronk of the University of Pennsylvania in 1940, and by Walter R. Hess of the 11 12 University of Zurich in the 1940s [9]. Hess won the Nobel Prize in 1949 for discovering the 13 14 functional organization of the diencephalon as coordinator of the activities of internal organs. 15 16 17 Karplus was married to Valerie von Lieben (1874-1938). The couple had four sons and 18 19 resided in the Palais Lieben-Auspitz, across the University’s main building. Karplus died on 20 21 22 11 February 1936 of a stroke [9]. He was interred at Döbling Cemetery. A manuscript on the 23 24 physiology of vegetative centers was published the following year in Bumke and Foerster’s 25 26 27 Handbook of Neurology [5]. 28 29 The eponym ‘Karplus sign’ signifies a modification in vocal resonance, or egophony, 30 31 whereby the vowel U spoken by the patient is heard as A on auscultation over a pleural 32 33 34 effusion [10]. 35 36 A street, Karplusgasse (48.17°N 16.34°E), was dedicated in 1956 at the boundaries of the 37 38 39 10th and 12th districts of Vienna, between the Meidling Trauma Center and the Kaiser-Franz- 40 41 Josef Hospital. 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 4 64 65

References 1 2 1. Anonymous (1963) Karplus, Johann Paul. In: Santifaller L, Obermayer-Marnach E (eds) 3 4 Österreichisches Biographisches Lexikon 1815-1950, Band III, Lieferung 13. Österreichische 5 6 Akademie der Wissenschaften, Wien, p 249 7 8 9 2. Karplus JP (1905) Über Familienähnlichkeiten an den Grosshirnfurchen des Menschen. Deuticke, 10 11 Leipzig 12 13 3. Karplus JP (1907) Zur Kenntnis der Variabilität und Vererbung am Zentralnervensystem des 14 15 Menschen und einiger Säugetiere. Deuticke, Leipzig 16 17 18 4. Karplus JP (1921) Variabilität und Vererbung am Zentralnervensystem des Menschen und einiger 19 20 Säugetiere: Familienuntersuchungen mit Berücksichtigung von Geschlecht und Entwicklung. 21 22 Deuticke, Leipzig 23 24 5. Karplus JP (1937) Die Physiologie der vegetativen Zentren (auf Grund experimenteller 25 26 Erfahrungen). In: Bumke O, Foerster O (eds) Handbuch der Neurologie, zweiter Band: 27 28 29 Allgemeine Neurologie II—Experimentelle Physiologie. Springer, , p 402-475 30 31 6. Kreuter A (1996) Deutschsprachige Neurologen und Psychiater. Saur, München, p 691-693 32 33 7. Marburg O (1936) Johann Paul Karplus, M.D. J Nerv Ment Dis 83:638-639 34 35 8. Marshall LH, Magoun HW (1998) Discoveries in the Human Brain: Neuroscience Prehistory, 36 37 38 Brain Structure, and Function. Humana Press, Totowa, NJ, p 245-246 39 40 9. Wang GH (1965) Johann Paul Karplus (1866-1936) and Alois Kreidl (1864-1928): two pioneers 41 42 in the study of central mechanisms of vegetative function. Bull Hist Med 39:529-539 43 44 10. White FA (2009) Physical Signs in Medicine and Surgery. Museum Press, Philadelphia, p 135 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 5 64 65

Fig. 1 Johann Paul Karplus. Photo by Max Schneider, 1908. From Julius Wagner-Jauregg’s 1 2 Lebenserinnerungen (Springer, 1950). 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Conflicts of interest None. 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 6 64 65 Figure 1 Click here to download Figure Figure 1 KARPLUS.tif