Acta Physiol 2011

A short history of physiology in the Netherlands

N. Westerhof
Departments of Physiology and Pulmonary Diseases, Institute for Cardiovascular Research (ICaR-VU), VU University Medical Center, Amsterdam, the Netherlands

Physiology is a branch of biology dealing with the functions and vital processes of living organisms, their parts and organs. In the later years of the Renaissance the dissection of human corpses, although still not permitted, became possible (Michelangelo, 14751564). The start of human anatomy can be considered to be the work of Andreas Vesalius, De humani corporis fabrica, (1543), the same year that Nikolaus Copernicus published his book on the revolution of the planets around the sun. The start of physiology is commonly considered to be in 1628, with the publication of Harveys (15781657) book: Exercitatio Anatomica de Motu Cordis et Sanguinis in Animalibus (An anatomical exercise on the motion of the heart and blood in living beings). This book became almost immediately known to the Dutch medical scientists. In other words, physiology in the Netherlands started soon after 1628, thus rst half of the 17th century. I will give a short description of what may be called the rst Golden age of physiology in the Netherlands. This period is followed by one age of consolidation. Then I will present a rather detailed description of the Second golden age of physiology. I will end with giving my views on present day physiology and its future. Thus, I will divide this review in the following sections (Fig. 1): 16281750 (Golden age of physiology) 17501850 (Consolidation) 18501975 (Second golden age of physiology) 1975present (Loss of Identity?)

Golden age of physiology 16301750

Very soon after the appearance of Harveys book, Dutch physiology started to blossom. The most important researchers in what we can now consider physiology deserve a short discussion. I will use the term physiologist loosely. In Figure 2 the most important physiologists of the period are depicted. Franz dele Boe (Franciscus Sylvius), was born in Hanau, now Germany, 15 March 1614 and died in Leiden 19 November 1672. He was a Dutch medical doctor, and professor at Leiden, as well as a scientist

(chemist, physiologist and anatomist). He was also an early supporter of R. Descartes, J.B. van Helmont and especially William Harvey. He understood Harveys work and theory well, and brought the concept of circulation of blood to the Netherlands. Sylvius was mentor of Johannes Walaeus, Niels Stensen, Jan Swammerdam and Reinier de Graaf. Jan de Wale (Johannes Walaeus, Koudekerke, Walcheren, 1604Leiden, 1649), studied philosophy and medicine in Leiden and later became professor at Leiden. He rst objected to Harveys theory, but Sylvius convinced him of its correctness. Walaeus is considered the founder of Dutch experimental physiology. He performed the so-called venous occlusion experiment to give support of the concept that blood returns to the heart, as part proof of the circulation. Niels Stensen (Nicolaus Steno, Kopenhagen, 10 January 1638Schwerin, 26 November 1686), was anatomist and physiologist and a student of Sylvius. He discovered the lymphatic system, and proposed that the heart is an ordinary muscle. He worked for a considerable time in the Netherlands and had contacts with the other physiologists, especially Jan Swammerdam. Although born form Lutheran parents, he later converted to the Roman-Catholic faith, and could therefore not be promoted to professor. He left the Netherlands and became Bishop in Germany (Kooijmans 2007). Jan Swammerdam (Amsterdam, 12 February 1637 Amsterdam, 17 February 1680) observed at an early age that muscle volume remained constant during a shortening contraction. He promoted the possibilities of the microscope consisting of two lenses, and was the rst to report on the function of red blood cells. He wrote a book on the respiration (Respiratione, 1667) and a book on insects that was re-published in 1737 with a foreword by Boerhaave and Gaubius. This book was written in Dutch: Alles in Hollandse, des Auteurs Moedertale (All in Dutch the authors mother tongue). This can be considered one of the rst signs that Latin as the scientic language was challenged. Reinier de Graaf (Schoonhoven, baptized 30 July 1641Delft, buried 17 August 1673) was a gynaecologist and gastrointestinal physiologist at Delft. He is

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Figure 1 Physiology in the Netherlands: main events.

best known for his studies of the function of pancreas in the living dog. His Roman-Catholic faith excluded him from a professorship. Anthonie van Leeuwenhoek (Delft, 24 October 1632Delft, 26 August 1723); was a so-called Physicist at Delft. Van Leeuwenhoek was educated as a bookkeeper and casher in Amsterdam. He started his rather lucrative drapery business in Delft in 1654. Having expertise in glass blowing, he made his own single-lens microscopes with a better resolution than the microscopes consisting of objective and ocular of the time as used by Swammerdam. Van Leeuwenhoek observed, among others, muscle bres, bacteria, spermatozoa and blood ow in capillaries and corresponded his ndings from 1674 through Sendtbrieven, to the Royal Society and was elected as a member of the Royal Society (London) in 1679/80. He died a rather rich man. Herman Boerhaave (Voorhout, 31 December 1668 Leiden, 23 September 1738) received his doctoral degree from the University of Harderwijk. He held,

for some time, three of the ve professorships in medicine at the University of Leiden. He was a botanist (director of the botanical garden at Leiden), chemist, humanist and a physician of world fame. He held the thesis that the pathology of an organ can only be understood from knowledge of its normal function. We can consider him as the founder of clinical teaching. His main achievement was to demonstrate the relation of symptoms to lesions. He had great interest in physiology, but was not a pure physiologist. However, as we will see that in the middle of 19th century physiology was initiated and begun by clinicians to give medicine a scientic basis, we mention him here. Jeroen Gaub (Hieronymus David Gaubius, Heidel berg, 1705Heidelberg 1780) studied, for personal reasons in the Netherlands at the University of Harderwijk, then went to Leiden, where he was a student of Boerhaave. He held the scientic proof that mind affects body and body affects mind. In 1758, his textbook of pathology (in Latin) appeared that

Figure 2 Physiologists of the rst Golden Age of Physiology in the Netherlands. 2011 The Authors Acta Physiologica 2011 Scandinavian Physiological Society, doi: 10.1111/j.1748-1716.2011.02286.x

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remained the standard text of pathology for almost 100 years. I believe the medical researchers of this Golden age of physiology can be called physiologists, even though their interests were many. The physiologists were very well informed about new theories and developments, and Harveys model of the circulation was readily accepted. In other words, these researchers had international contacts and also knew each other. Most of them travelled, three were foreign born. Van Leeuwenhoek may not have travelled much but his international contacts were very good, especially his membership of the Royal Society of London. By the end of the 18th century, the scientic language changed from Latin to Dutch.

diseases are based on the cellular processes. This concept helped to convince the medical profession that physiology, as it was perceived then, forms the basis of (internal) medicine and should be introduced in the medical curriculum.

Second golden age of physiology 18501975 New names

By the end of 18th century, some need existed to intensify communication and exchange. This resulted in the foundation of a number of Genootschappen (Societies). One of them is the still existing Genootschap ter bevordering van de Geneeskunde (Society for the promotion of Medine) that later was extended to Genootschap ter bevordering van Natuur-, Genees- en Heelkunde. However, in the middle of the 19th century, a real strong revival of the sciences (Charles Darwin, 1859, the Origin of Species; Dmitri Mendelejev, 1869, the periodic table) and the strong growth of technology (steam locomotives and railroads, rst steel steamship crossed the Atlantic, 1852) took place all over Europe. Several specialized branches of the medical sciences arose: gastrointestinal physiology (William Beaumont, 17851853); pathology and cell physiology (Rudolf Virchow, 18201902); bacteriology (Louis Pasteur, 18221895, Ignaz Semmelweis, 18181865 and Robert Koch, 18431910). In integrative physiology, it is Claude Bernard (1813 1878) who can be considered the rst true general physiologist, but others were prominent too: Ivan Pavlov, 18491936 psychophysiology; Charles Sherington, 18571952 (neurophysiology), Carl F.W. Ludwig (18161895) and Ernest Starling, 18661927 (cardiovascular physiology and hormones), Henry P. Bowditch 18401911 (1871, rst physiological laboratory for students in the USA).

Consolidation 17501850
After 1750 a standstill followed. This so-called Consolidation period lasted for about 1100 years (Lindeboom 1981). It was not a local phenomenon but was generally seen in Europe in the (medical) sciences. It was especially clear in the Netherlands as can be seen from the following examples. While the use of thermometer in the clinic was introduced by Anton de Haen, a pupil of Boerhaave in the middle of 18th century in Vienna, and used in daily clinical practice, this instrument was not used in Utrecht until 100 years later. Also in the middle of 19th century, the microscope was not introduced to medical students in Leiden. Teaching in physiology consisted of reading Latin texts (Gaubius textbook was still the standard); also auscultation (invented by Auenbrugger and introduced in the clinic by Laennec in 1819) was not or hardly used in 1850. The origin of standstill of medical sciences seems to be the result of rise in philosophical idealism in Germany, mainly under the inuence of I. Kants (17241804) and F.J.W. von Schellings (17751854) Natur Philosophie: the intelligent human is able to understand the natural phenomena; experimental, science-based approach, is not necessary (Lindeboom 1981). Around 1840, a number of young medical doctors in Germany became proponents of the concept medicine = science and proposed that science should be introduced into the medical curriculum. This ended the romantic philosophical period, in part also thanks to Rudolf Virchow (18201902), founder of cellular pathology. Especially his text Die Cellularpathologie in ihrer Begrundung auf physiologische und patholog ische Gewebelehre (Cellular pathology as the basis of physiological and pathological histology) of 1858, where the cell was introduced as the smallest unit of the organism, made it convincingly clear that most

New journals
In the middle of the 19th century also many new journals of physiology originated. In Germany in 1835, the Archiv fur Anatomie, Physiologie und wissenschaf tliche Medicin (until 1876) was founded. In 1842 and 1844, the Archiv fur physiologischen Medicin and the Zeitschrift fur rationelle Medicin followed. In 1847, the Archiv fur pathogische Anatomie und Physiologie und fur klinische Medicin, was founded by R. Virchow and B. Reinhardt (now called Virchows Archiv). In 1854, Albrecht von Graefes Archiv fur Ophthalmolo gie (now called Graefes Archive for Clinical and Experimental Ophthalmology) was founded; and in 1862 Fresenius Journal of Analytical Chemistry,

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originally called Zeitschrift fur analytische Chemie started. Pugers Archiv (now Pugers Archiv Euro pean Journal of Physiology) started in 1868, while the Journal of Physiology (London) was established in 1878; and somewhat later, in 1898 the American Journal of Physiology was begun. The Journal de Physiologie (Paris) began in 1906. At present there exist more than 120 journals with the term physiology in their title. The Nederlands Tijdschrift voor Geneeskunde (the Netherlands Journal of Medicine, NTvG), founded in 1857, played an important role in the communication between physiologists, and between physiologists and clinicians in the second half of the 19th and early 20th century. Table 1 (Heteren van 1991) gives an overview of the (retrievable) publications of the professors of physiology between 1850 and 1950 of the four original medical schools. It may be seen that the NTvG was used for more general information as well as for publication of research articles. The sections in the journal, next to Research, namely News, Opinion, Clinical practice, Perspectives and Varia, contain many subjects of more general interest on which the physiologists had an opinion. The interest of the physiologists in general aspects of society, medicine and

education with respect to research is also made quantitative in Table 1. Most physiologists published their research articles in German journals. Einthoven published only six articles on research in the NTvG, but 33 articles in international journals. It should also be noted that most research articles of the physiologists, quite contrary to present day articles, were singleauthored.

The Netherlands makes up its arrears

The international change affected the Netherlands as well, where it was given extra impetus after the, (un?) founded, publication by Moleschott, (then at Heidelberg, originally from Leiden, later in Italy) that medical sciences in the Netherlands were running behind compared with other European countries. The Dutch originator of the change appears to be J.L.C. Schroeder van der Kolk (17971862) with his work in neurology and experimental physiology, but the real driving force of the developments in physiology was F.C. Donders in Utrecht. The law of 1868 by J.R. Thorbecke ended the complex system of diplomas and introduced a single, country-wide, diploma to practice medicine in the Netherlands.

Table 1 Publications of Professors of Physiology at the Dutch universities; second half 19th and rst half 20th century NTvG (res) Utrecht University FC Donders 13(7) ThWEngelmann 7(2) H Zwaardemaker 326(45) AKM Noyons 5(2) J Jongbloed 32(10) Groningen University I van Deen 20(10) D Huizinga 0 HJ Hamburger 59(33) FJJ Buytendijk 20(3) University of Amsterdam A Heynsius 51(14) W. Kuhne (1) T Place 5(0) GA van Rijnberk 787(14) Leiden University A Heynsius 51(14) W Einthoven 38(6) GGJ Rademaker 12(7) JW Duyff 26(3) EJP Fres Grae Virch Ergeb Eng JoP + AJP Other TOT

5 48 12 3 3

25 1 1 9 1 1 6 2+1 1

3 2 15 1

40 52 86 10 20 10 11 51 2 25 32 1 16 25 39 9 6

2 6

1 10

2 2

9 1

2 3

20

2 1

4 1

9 29 1

2 1 1 2 0+1 1 2

NTvG, Nederlands Tijdschrift voor Geneeskunde (total number with number of research articles in parentheses; EJP, Pugers Archiv, European J Physiology; Fres, Fresenius Journal of Analytical Chemistry; Grea, Graefes Archive for Clinical and Experimental Ophthalmology; Virch, Virchows Archiv; Ergeb, Ergebnisse der Physiologie, biologischen Chemie und experimentellen Pharmakologie; JoP, J of Physiol (Lond); AJP, American J of Physiology. The information on EJP, Fres, Grea, Virch, Erg, and other is obtained from Springers Archives.

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These changes, together with the wish for scientic underpinning of medicine implied that physiology became mandatory in the medical curriculum, and that the physiology as basic medical science needed laboratory facilities.

Physiologys second growth

After the Second World War, we see several changes taking place in the universities that affected physiology. Four new medical schools were begun, each with a department of physiology. In 1950, the Vrije University of Amsterdam founded a medical school, after the unsuccessful rst start with F.J.J. Buytendijk in 1919 who, in 1925 became professor of physiology in Groningen. The rst professor of physiology at the Vrije Universiteit was A.Th. Knoppers, who left after only 2 years for industry, and was succeeded by A.W.J.H. Hoitink in 1953. The (now) Radboud University in Nijmegen started with a medical school including physiology in 1951, where F.J.A. Kreuzer was the rst professor. The Erasmus University in Rotterdam began with a medical school in 1966, where M.W. van Hof, was the rst professor. Maastricht University founded in 1974, set-off with a medical school with R.S. Reneman as the rst professor of physiology. In summary, by 1974 there were eight universities with medical schools in the Netherlands, each with its department of physiology.

Physiology develops rapidly from 1848

Between 1848 and 1866, all four Universities in the Netherlands that offered a medical curriculum, nominated a professor of physiology and built a research laboratory. The rst professors of physiology at the Dutch Universities were: F.C. Donders (Utrecht, 1848 1869), I. (sometimes J.) van Deen (Groningen, 1851 1869), A. Heynsius (Amsterdam, 18581868). Heynsius later moved to Leiden and was there the rst professor of physiology (18661885). A complete overview of the professors (chairpersons) of physiology from 1858 to present is given in Table 2. The research performed was very closely related to clinical problems. The clearest example is, of course, Donders at Utrecht who combined, as professor of internal medicine, his clinical duties with his professorship in physiology. It is also interesting to note that the early professors of physiology published on many subjects, usually as single author, and often in the form of rather short publications. Professors of physiology were also involved in more public affairs. Examples are D. Huizingas book: Schetsen uit het leven (Images from life); and books for grade school pupils; Hamburger was vice-president of a society to expand permission to enter university for graduates of the secondary school HBS and not only students with a gymnasium (latin school) diploma (Gemert van et al. 1989, Westerhof 2010, WWW document). Many physiologists of the rst century of organized Dutch physiology (18501950) were member of the editorial board of NTvG; A. Heynsius, I. van Deen, W. Einthoven and especially G.A. van Rijnberk who was editor in chief, then called gerant, from 1913 to 1946 (Heteren van 1991, Quak 2005). Figure 3 shows the portraits of the professors (chairs) of physiology until 1950. Teaching. Most teaching of physiology to medical students was in a format of classroom lectures with occasional demonstrations. It was van Rijnberk who, with his appointment in 1909, was the rst to organize laboratory teaching, including animal experiments. This was an international trend with Henry P. Bowditch as the rst in the USA. Almost all professors wrote or contributed to textbooks of physiology, most of which in Dutch. An example is van Rijnberks textbook of physiology in the 1930s and 1940s entitled: Neederlandsch Leerboek der Physiologie, which appeared in seven volumes.

Physiological research in the Dutch Universities

Utrecht University. The physiology in Utrecht, starting with F.C. Donders, was oriented to the sensory system. Originally, vision was the main theme. Donders was a giant in many aspects and had broad interests; he became most famous for his work on vision. Donders was followed by W.Th. Engelmann who was mainly interested in neuromuscular physiology. Engelmann returned to Germany in 1897, and his successor was H. Zwaardemaker who worked on sensory physiology of smell and taste and on phonetics. Both had other interests too: Engelmann studied the isolated heart, and Zwaardemaker studied the biological meaning of radiation with Professor L.P.H. Eijkman, a brother of C. Eijkman who received the Nobel Prize of Medicine or Physiology. His theory of radiation in relation to impulse formation of the heart was shown to be incorrect by Yngve Zotterman. I mention here two physiologists of great fame. Rudolph Magnus (Braunschweig, Germany 1873Switzerland 1927) was professor of pharmacology at Utrecht University with interest in neuropharmacology and neurophysiology, from 1908 until his sudden death in 1927. Rudolf Magnus worked in Liverpool in 1908 on the physiology of stature-movement and muscle tension. This physiological work made him world famous, and, as was learned 50 years later, it formed the basis of his nomination for the Nobel Prize of 1927. Unfortunately, he died suddenly in the summer of that year. The Rudolf Magnus Institute of Neurosciences is named after him. Herman J. Jordan (Parijs 1877Zeist

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6
University of Amsterdam A Heynsius 1858 1868 W Kuhne 1869 1871 T Place 1871 1909 GA v Rijnberk 1909 1946 J ten Cate 1946 1957 JThF Boeles / LN Bouman GJ Tammeling 1953 1985/1971 1985 1971 1980 LN Bouman 1985 1997 JH Ravesloot 2001 present 2009 Part Embr &Anat 2002 Clinical Sciences WJ Rietveld 1980 2000 AA Knoop 1969 1994 N Westerhof 1994 1996 GJ Tangelder 1996 present AA Verveen 1969 1970 ATh Knoppers 1950 1952 FJA Kreuzer 1951 1985 MW van Hof 1966 1993 CH van Os / B Oeseburg H Collewijn 1989 2007 / 1989 1999 1993 2002 RJM Bindels 2007 present RS Reneman 1974 1990 2002 MA Allessie Dept Neuroscience 1990 2004 MA Post 2004 present JW Duyff 1946 1969 GGJ Rademaker 1927 1946 FJJ Buytendijk 1919 1924 W Einthoven 1885 1927 A Heynsius 1866 1885 Leiden University VU Amsterdam Radboud University Erasmus University Maastricht University AWJH Hoitink JA Bernards 1953 1969 1985 1988

Table 2 Professors of Physiology at the Dutch Universities (Ordinarius/Chairman only)

Physiology in the Netherlands

Utrecht University Groningen University

FC Donders 1848/54 1888

I van Deen 1851 1869

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ThW Engelmann 1889 1897

D Huizinga 1870 1901

H Zwaardemaker HJ Hamburger 1897 1927 1901 1924

AKM Noyons 1928 1941

FJJ Buytendijk / R Brinkman 1925 1946 / 1933 1956

J Jongbloed 1942 1965

HD Bouman 1947 1948

PA Biersteker 1966 1988

MJN Dirken 1949 1956

WL Mosterd 1988 1993

R Brinkman 1956 1960

HJ Jongsma 1993 2002

WG Walter / WG Zijlstra 1960 1977/1960-1990

MA Vos 2003 present

C Rashbass 1977 1982

D Kernell 1997 2000

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HWGM Boddeke 1998 pres. Dept Neuroscience Acta Physiol 2011

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Figure 3 Professors, chairs, of Departments of Physiology between 1850 and 1945.

1943) was educated in Germany and received his PhD degree in Bonn, Germany (Jong de 2002). He attended the 9th International Congress of Physiology in Groningen in 1913 and, through his contact with Magnus, was appointed at Utrecht University, where he became professor in 1915 and full professor in 1918 in physiology/zoology in the school veterinary medicine, and founded an institute (Zoologisch Station Den Helder). He was forced to retire in 1942 in the middle of the Second World War and died in 1943. A.K.M. Noyons, rst continued the research line started by Zwaardemaker but, with his successor, J. Jongbloed, got interested in the effects of acceleration on the body. Jongbloed was very interested in space and sports physiology, but also played an essential role in the development of rst heart-lung machine. W.L. Mosterd and P.A. Biersteker again were interested in

sports physiology, but research in cardiac physiology also took place. H.J. Jongsma shifted the direction to the electrophysiology of the (cardiac) cell. His most important contribution was the electrical interaction of cardiac cells through gap junctions. Integrated cardiac electrophysiology from cell to organism is the research theme at present under the leadership of M.A. Vos. University Groningen. I. van Deen was born in Germany, and educated at Copenhagen University, Denmark. He assumed de name Deen, and performed research neuro-physiology. His successor D. Huizinga was interested in blood and its components, and also in Abiogenesis (the study of how life on earth could have arisen from inanimate matter). Subsequently, H.J. Hamburger became professor (Gemert van et al. 1989). He obtained a degree in chemistry followed by

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a degree in medicine under Donders. His general concept was that physico-chemical principles play an important role in physiology and thereby also in medicine. In 1913, he organized the 9th International Congress of Physiology at Groningen, the predecessor of the IUPS meetings (Westerhof 2010, WWW document). F.J.J. Buytendijk after his medical study, changed his interests, via biology and then clinical neurology, to psycho-physiology with the emphasis on psychology. R. Brinkman chaired a section of physiological chemistry from 1933 onwards. This group became an independent laboratory with Brinkman as chairman. While Brinkman concentrated on chemical physiology H.D. Bouman was appointed in physiology and after only 2 years he was succeeded by M.N.J. Dirken. In the following interim period between 1956 and 1960 R. Brinkman was chairman of physiology and in 1960 left to chair the department of Radiopathology. In 1960, the department was split into two, and W.G. Walter and W.G. Zijlstra were appointed as chairs of the two departments of physiology (Walter 1961). Walters research was in neuromuscular physiology and Zijlstras work was in organ physiology, with emphasis on blood chemistry and oxygen transport. In 1977, Walter was succeeded by C. Rashbass who stayed till 1982. Zijlstra retired in 1990. After an undetermined period, physiology consolidated into a single laboratory of medical physiology with Kernell as chairman. After Kernell left, the physiology as department was dissolved. An institute of neurosciences was formed, where H.W.G.M. Boddeke, since 1998, is responsible for physiology. J.A. Bernards and L.N. Bouman the authors of the textbook Medische Fysiologie were, in later editions (2008), joined by Boddeke. University of Amsterdam. The physiology in Amsterdam began with physiological chemistry of blood, the main interest of A. Heynsius. Later Heynsius also published about the circulation and respiration. Heynsius left after 10 years for Leiden and W. Kuhne succeeded him. Kuhne was born in Hamburg, now Germany, and studied with all great physiologists of the time: Emil du Bois-Reymond at Berlin, Claude Bernard in Paris, and Carl F.W. Ludwig in Leipzig, now Germany and Ernst W. von Brucke in Vienna, Austria. Kuhne was a top researcher working mainly in chemical physiology (of blood) and he coined the word enzyme. Also nerve-muscle physiology was his interest. T. Place, his successor, published only a few articles on the nervous system. His successor was G.A. van Rijnberk, a very colourful person, was educated in Italy, and had an immense knowledge of art. He was a great teacher both feared and adored by the students but was more a teacher and organizer than a

researcher. He published only a few research articles but edited a textbook of physiology. (see above). He was editor in chief of the Nederlands Tijdschrift voor Geneeskunde from 1913 to 1946, and his editorial comments were many (hundreds). When he celebrated his 25th anniversary of his professorship, an overview of his work was published in the book entitled: Een greep uit het werk van van Rijnberk (1934). His successor, J. ten Cate, who was born in St. Petersburg, Russia, and a descendant of Dutch merchants that moved to Russia in the 18th century. He studied medicine in Moscow, Berlin and Paris. He worked with Pavlov in St Petersburg before coming to Amsterdam, which inuenced his research interests: conditional reexes and electroencephalography are keywords of his research. J.Th.F. Boeles and L.N. Bouman were, interchangeably, chairman of the department of physiology. Boeles studied nerve action in the broadest sense, from motor nerves to sympathetic nervous effects on the heart. Boumans main research was the electrophysiology of the heart, with emphasis on the pacemakers and the conduction system. Bernards and Bouman wrote the textbook Fysiologie van de mens. In later editions, Boddeke joined them and the title changed to the present one: Bouman, Bernards en Boddeke, Medische Fysiologie (2008). After the retirement of Boeles and then of Bouman, the position remained open for a few years, but in 2001 H.J. Ravesloot lled the position of professor of physiology. His main interest was cellular physiology. In 2009, the department of physiology merged with the department of anatomy and embryology to form the department of Anatomie, Embryology and Physiology. Leiden University. A. Heynsius, became the rst professor of physiology in Leiden, after having this position for 10 years in Amsterdam. Physiological chemistry of the blood and its constituents was his primary interest and later he also published about the circulation and respiration. His successor was W. Einthoven, who defended his doctoral thesis in Utrecht under the supervision of F.C. Donders, and became full professor of physiology at the age of 25 in 1885. Einthoven was most famous through his work on electrocardiography for which he received the Nobel prize in 1925. The study of electrocardiography required physical and technical expertise, which Einthoven picked up during his medical studies when he worked with physicists like C.D.H. Buys Ballot and others. Next to the electrical phenomena of the heart he was interested in neurophysiology, especially the inuence of the brain on body position and movement. Einthoven was succeeded by G.G.J. Rademaker, neurophysiologist and neurologist, who was a pupil of Rudolf Magnus (Braunschweig, Germany 1873Switzerland

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1927). Magnus was professor of pharmacology at Utrecht University with interest in neuropharmacology and neurophysiology, from 1908 until his sudden death in 1927. The Rudolf Magnus Institute of Neurosciences is named after him. He wrote textbooks in German, French and English (sensory system, vision, hearing and vestibular function). In 1946, he left physiology for neurology. Between 1946 and 1969, J.W. Duyff was the professor of physiology at Leiden. He played an important role in the resistance movement during the Second World War. He was a very good organizer, and was editor in chief of Acta Physiologica et Pharmacologica Neerlandica from 1950 to 1969. He was (vice-) editor of the Nederlands Tijdschrift voor Geneeskunde (NTvG) from 1956 to his sudden death in 1969. His main interests were vision and audition and his articles were mostly published in Acta Physiologica et Pharmacologica Neerlandica and the NTvG. He also wrote a short review on Dutch physiology (Duyff 1952). Professor Duyff as chairman of Physiology organized the now appropriately called 22nd IUPS meeting in Leiden in 1962. Duyff was succeeded by G.J. Tammeling, and under his chairmanship four groups were formed: biological chance processes; chronobiology and the adaptation to the external milieu; respiratory physiology and physiological physics. W.J. Rietveld who was Tammelings successor was interested in and published many articles on chronobiology. The physiology became part of a department of clinical sciences in 2000. VU University Amsterdam (formerly Vrije Universiteit of Amsterdam). After the rst attempt to start a school of medicine (19191924), the medical school at the Vrije Universiteit (Now called VU University) started in 1950. A.Th. Knoppers was the rst professor of physiology, but left for industry in 1952. His main interest was pharmacology. A.W.J.H. Hoitink succeeded him in 1953; he was a physiologist with very broad interests and recently retired from Batavia (Indonesia). One of his research subjects was ballistocardiography, then a newly proposed technique (Jongbloed 1951) given a physical basis in Utrecht by H.C. Burger and A. Noordergraaf. A.A. Knoop, his successor began in 1969 and initially continued this research direction. However, ballistocardiography was too complex for interpretation and clinical application. It was therefore decided to concentrate the entire laboratory on cardiovascular physiology, with strong emphasis on haemodynamics. After a bridging period of 2 years by N. Westerhof, G.J. Tangelder, member of the department of physiology at Maastricht University, became chairman of physiology. Presently, the department covers cardiovascular physiology from molecule to organism.

Radboud University Nijmegen (formerly Catholic University Nijmegen). With the foundation of the school of medicine in, what is now called Radboud University, professor F.J.A. Kreuzer was chosen as head of the department of physiology. Kreuzer came from Switzerland, worked in the departments of physiology at Fribourg, Switzerland, Shefeld, UK and Dartmouth NH USA, before coming to Radboud University. His main interest was oxygen transport by blood and diffusion, and he developed oxygen electrodes. J.A. Bernards treaded in the steps of Kreuzer and worked mainly on the effects of pulmonary gas embolism on circulation and respiration. Bernards, with Bouman, and later Boddeke are the authors of the textbook Medische Fysiologie (Bouman et al. 2008). Bernards was followed by C.H. van Os and his interest was the physiology of water and salt management and cell biology of the kidney. R.J.M. Bindels succeeded van Os and carries out research on transport physiology in epithelia of the kidney and intestine, especially the molecular and cellular physiology of electrolyte transport. Presently, the department of physiology exists of three research groups: integrative physiology, ion transport and osmoregulation. Erasmus University Rotterdam. The department of physiology started in 1966 with Professor M.W. van Hof as chairman. Van Hof studied the physiology of brain function in the widest sense. He retired in 1993 and his successor was H. Collewijn who mainly studied vision and the vestibular system. In 2002, the department was dissolved and became part of Neurosciences. Maastricht University. In 1974 with the start of new university at Maastricht, R.S. Reneman founded the physiology, and was soon joined by F.I.M. Bonke. Reneman was educated as an anaesthesiologist in Utrecht and worked several years with Janssen Pharmaceutica before joining Maastricht. The research was concentrated on the cardiovascular system, namely, electrophysiology of the heart, cardiac and large artery function, and with special attention to the microcirculation. Also the development of ultrasound techniques should be mentioned. M.A. Allessie, followed Reneman in 1990, and he studied electrophysiology of the heart with special emphasis on atrial brillation. Presently, M.A. Post is chairman of the department and his main research interests are angiogenesis/collateral formation and vascular healing and remodelling after arterial injury.

Societies and meetings

International. Following the decision of a group of physiologists in 1885 regular, international, meetings of

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physiology started with the 1st International Congress of Physiology at Basle, Switzerland, in 1889. These meetings were held every third year (excepting 1916, 1941 and 1944), and physiologists from Europe and the Americas participated. The last, 18th meeting, was held in 1950 in Copenhagen Denmark. Subsequently, the international meetings were organized by the International Union of Physiological Scientists. These meetings, being a continuation of the international congresses started in 1953, with the rst IUPS meeting counted as the 19th congress. After 1989 the IUPS meetings were held every 4th instead of every third year. The 37th meeting will be in Birmingham, England in 2013. In 1913, Hamburger organized the 9th International Congress of Physiology at Groningen (Westerhof 2010, WWW document). The number of participants was 594 and 150 presentations (26 from the Netherlands) were given. The prize for the best Dutch doctoral dissertation is named after Hamburger and is awarded yearly since 1987, by the Netherlands Society of Physiology. Among the presentations of this meeting was one by J.G. Dusser de Barenne (18851940). Dusser de Barenne studied medicine in Amsterdam, worked and was lecturer at Utrecht University and went to Yale (New Haven, CT, USA) in 1930 to set up a laboratory of neurophysiology. He died of coronary disease in 1940. The oeuvre-award of the Netherlands Society of Physiology is named after Dusser de Barenne. Professor J.W. Duyff chairman of Physiology at Leiden organized the, now appropriately called 22nd IUPS meeting in Leiden in 1962. This was a very large international congress. Europe. As the forming of FEPS, the Federation of European Physiological Societies (Federation European Societies of Physiology, FEPS 1991, WWW document), there are also meetings for mainly European physiologists. Presently, the physiological societies of 31 countries (counting England and Ireland as 1) contribute to the FEPS. The rst three meetings were in Maastricht (1995), Prague (1999) and Nice (2003). Subsequently, so-called joint meetings were organized by a combination of host country and the FEPS. Examples are the joint meeting of the Physiological Society and FEPS in Bristol, UK (2005); the German Physiological Society and FEPS, Munich Germany (2006); the Slovak Physiological Society and FEPS, Bratislawa, Slovakia (2007); the Slovenian and Austrian Physiological Societies and FEPS, Ljubljana Slovenia (2009). The following joint meetings are planned: the German-Scandinavian Physiological Societies and FEPS, Copenhagen, Denmark (2010); the Turkish Physiological Society and FEPS, Istanbul, Turkey (2011); the Spanish Physiological Society and FEPS, Santiago de Compostela, Spain

(2012); the Hungarian Physiological Society and FEPS, Budapest, Hungary (2013). The journal of the FEPS is Acta Physiologica, formerly the journal of the Scandinavian Physiological Society: Acta Physiologica Scandinavica. National. The rst, founding, meeting of the Nederlandse Vereeniging voor Physiologie en Pharmacologie (Netherlands Society of Physiology and Pharmacology) was held April 15, 1916 (van Rijnberk 1916). It was the rst Day of Physiology, a whole day (Saturday) with about 100 participants: physiologists, practicing physicians and medical students. The start of this society shows a denitive break with anatomy. About 1950 the societys name changed from Nederlandse Vereniging voor Physiologie en Pharmacologie to Nederlandse Vereniging voor Fysiologie en Farmacologie, to adhere to the changed Dutch spelling rules. In 1975, the Nederlandse Vereniging voor Fysiologie en Farmacologie decided to rearrange their organization by splitting into two sections: Physiology and Pharmacology, but continuing as a single society. However, nice an idea, in 1979 an ofcial divorce followed, and from then on physiologists were united in the Nederlandse Vereniging voor Fysiologie. National meetings. The members of the Nederlandse Vereeniging voor Physiologie en Pharmacologie convened several times per year up to about 1960. The meetings were announced in the Nederlands Tijdschrift voor Geneeskunde, and the location rotated between the universities. Occasionally, a foreign sister society was invited. For instance, in 1925 a meeting was held in Leiden, with members of the Physiological Society of the UK. That meeting included a visit to Amsterdam. In 1958, the meeting was held in Amsterdam together with the Belgisch Genootschap voor Fysiologie (Belgian Society of Physiology), and in 1975 there was a combined meeting in Gent with the Belgisch Genootschap voor Fysiologie. In 1959, the Nederlandse Federatie van Medisch en Biologische Verenigingen (Dutch Federation of Medical and Biological Societies) started to organize meetings broader than physiology alone, mainly animal biology, in the Netherlands, and many physiologists participated in these meetings. However, it was felt that a regular exchange of developments in physiology was more useful, and in 1973 yearly meetings of the Nederlandse Vereniging voor Fysiologie began.

Papendal symposia
The rst so-called Papendal-meeting was organized in 1973, they were fall meetings. This meeting evolved, in 1984, into a 2-day symposium of physiology, without

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pharmacology and on a regular, yearly, basis. The later started Young Physiologists Day (1997) was originally a separate meeting in the spring, where the young physiologists could exchange ideas with presentations and posters. In 2004, this Young Physiologists Day was incorporated in the Papendal Symposium. Thus, from 2004 the Papendal Symposium is a 2-day symposium, in early December, with the rst day devoted to the Young Physiologists, and in the evening the lecture of the Hamburger Prize recipient. The second day is on a general aspect of physiology with Dutch and foreign expert speakers. While the location, the conference centre Papendal, will be changed the name of the yearly meeting will remain as known as Papendal-meeting.

See for example the article by Engelmann (1870). This argumentative structure was gradually abandoned in the early 20th century. Now, the generally accepted format of articles appeared in the American Journal of Physiology in the early part of the 20th century and it became standard after in the 1930s. In the Netherlands, this format was then not commonly accepted yet. See for instance the article by Duyff (1946) and compare it with a modern article. However, in the 1950s this standard form of articles became common in the Netherlands as well. Dutch journals. With the start of organized physiology in 1916 (van Rijnberk 1916), it was still generally felt that publications had to appear in Dutch contexts. The Archives Neerlandaises des Sciences Exactes et Natur elles, published by la Societe Hollandaise des Sciences a ` Harlem started in 1866 and lasted until 1911. In 1918, a successor journal was started, entitled: Archives Neerlandaises de Physiologie de lHomme et des Animaux. In 1927 Archives Neerlandaises de Phonetique Experimentale (19271947) split off, and in 1931 another journal appeared entitled Acta Brevia Neerlandica de Physiologia, Pharmacologia, Microbiologia (in short Acta Brevia). It was a merger of Archives Neerlandaises de Physiologie and Acta Brevia that formed, in 1950, the Acta Physiologica et Pharmacologica Neerlandica. This new journal, with J.W. Duyff as editor in chief, was very modern in layout. It appeared four times a year, the languages were English, German and French, but most articles were in English. Abstracts were presented in three languages. The journal also published abstracts of the meetings of the Nederlandse Vereniging voor Fysiologie en Farmacologie, and can be considered as the journal of the Netherlands Society of Physiology. By 1968, it became clear that Dutch researchers were more and more preparing their articles for international English-language journals, and that Acta Physiologica et Pharmacologica Neerlandica, having a rather small number of subscribers, could not attract enough articles. Meanwhile Pugers Archiv was exploring chang ing into a European Journal of Physiology. In a letter to the editors of Pugers Archiv by Boeles as chairman of the Physiologists Convent, it was mentioned that Acta Physiologica et Pharmacologica Neerlandica was in der Agone (in agony). However, the Dutch Physiologists Convent was hesitant and no countries supported this idea of making a European Journal of Physiology, and a new European journal was not formed. In January 1975, Pugers Archiv was indeed transformed into Pugers Archiv European Journal of Physiology (First volume 361). The last volume (number 15) of the Acta Physiologica et Pharmacologica Neerlandica appeared in 1969, and the journal was absorbed by European

Physiologists convent
In 1961, the question arose how (some) medical students who wished to be a physiologist should be educated. This idea was based on a similar action of the pharmacologists. Also there was a proposal to organize a country-wide rst year of the medical curriculum. To discuss these matters and other, more general aspects, such as nominations of new Professors of physiology in the Netherlands, professor Boeles chairman of physiology at the university of Amsterdam and on suggestion of Prof. J.W. Duyff at Leiden, in 1966, asked all colleagues (seven at the time, and eight after 1974) to form the Physiologists Convent. To keep the number of participants small it was decided to let one representative per university participate. It turned out that this was almost always the department chairman. The convent started to make an inventory of the physiology courses in the different universities, but found that the physiology curriculum differed too much to come to a core curriculum. Other subjects, such as the problems with Acta Physiologica et Pharmacologica Neerlandica were also discussed (see below). After the 1980s the meetings were held less and less frequently. The impression is that the convent was not sufciently united and did not sufciently emphasize the importance of physiology in research and, especially teaching.

Publications
Writing format of articles. Up till the Second World War most scientic publications of the Dutch physiologists were in German journals and only few were in French and English journals, Also a substantial number of publications appeared in the Nederlands Tijdschrift voor Geneeskunde. Originally scientic articles were based on a format, where the authors thought processes that gave rise to the results were presented and defended in the order in which the experiments were carried out (Suppe 1998).

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journal of Pharmacology in the same year. The titles of the articles in Acta Physiologica et Pharmacologica Neerlandica can be found in PUBMED, but the abstracts and full articles of the journal are not electronically available. However, paper versions are obtainable in the university libraries of Leiden and VUmc library. Between 1907 and 1957, the Dutchlanguage journal Tijdschrift was in existence. The editor was van Rijnberk and the journal was mainly meant for the broad general public. The Nederlands Tijdschrift voor Geneeskunde still exists but is presently little used by physiologists for publication.

Loss of identity?
Professors of physiology performed not only teaching and research but had also societal interest (see Table 1). They mainly published in the Nederlands Tijdschrift voor Geneeskunde, and not only research articles but also notes and papers on general subjects. Also physiologists regularly published in newspapers about developments in medicine, especially in the late 19th and early 20th century. A later example is J. Jongbloed who wrote regularly in Utrechts Nieuwsblad (the daily newspaper of Utrecht) about space physiology. Furthermore, professors of physiology worked in general very closely with clinical researchers, this was how physiology was initially started, namely to give medicine a scientic basis. After Second World War, especially in the 1960s and 1970s the number of (medical) students increased enormously. Also societal relevance of the medical research became more and more important. In the same period, research in the medical sciences became divided in a number of specialized areas. This was, in part, the result of the great successes in cell biology, molecular biology, and somewhat later genomics and protein biology. In 1977, the American Physiological Society decided to split the American Journal of Physiology into a number of sections. This decision was unavoidable as specialization of researchers made it impossible for them to read all news in physiological research. Nevertheless, these developments resulted in decrease in recognition of physiology as the science of functions and vital processes of living organisms, their parts and organs. From the 1980s researchers, including professors where evaluated on the basis of research articles (especially citations), leaving, next to teaching, and the increased administrative duties, even less time for other activities. For example, J.W. Duyff was the last physiologist who was (vice)-editor in chief of the Nederlands Tijdschrift voor Geneeskunde. Also the Physiologists Convent did not propose a united country-wide programme for teaching and research had grown too far apart from direct clinical

applications, it became almost a separate discipline (too little translational research). These observations may have led to the idea that there is no need for separate departments of physiology. At present the translation of the enormous amount of knowledge gathered in sub-branches of biology such the cell, molecule and gene is not sufciently studied in the context of the function of the organ or organism as a whole. At the cellular level systems biology was started about 1990, where an interaction of many complex mechanisms is combined to understand overall cellular function. This approach requires the use and construction of computer models (Alberghina & Westerhoff 2008). Also, recently NOW the Nederlandse Organisatie voor Wetenschappelijk Onderzoek (NWO) with the programme Centres for Systems Biology Research stimulates and funds research from molecule to organism. Luckily physiology at the organ and organism level is moving in that direction as well. To better understand the physiology the step upwards to the organ or organism is made through the so-called physio(no)me (Bassingthwaighte 2000).

Future
Research. In his review, C.A. Pekelharing (18481922, professor of physiology, comparative anatomy and pathology at Utrecht University) in 1907 made it clear that the good medicine is based on the scientic approach (Pekelharing 1907). From the 19th to far in the 20th century physiology was the leading basic science and was integrated with the clinical sciences. For example, Alfred Noble created the Nobel Prize in Physiology and Medicine rather than a prize in medicine. The specialization in the research, i.e. to cell, molecule and gene, moved physiology further away from direct clinical practice. However, it is becoming more and more clear the translation of this basic knowledge to the organ an organism level is necessary. This is not only seen in the IUPS Physiome Project (IUPS), WWW document but also recognized by NWO (the Netherlands Organization for Scientic Research) with its NWO-programme complexity. At the physiological level this is materializing through the IUPS Physiome Project (IUPS), WWW document, a worldwide public domain effort to provide a computational framework for understanding human and other eukaryotic physiology. A related approach is Computational Physiology (Hunter & Nielsen 2005), from Genome to Physiome or Physionome physio (life) + ome (as a whole). In short, it appears that collaboration and computers are key words with good translational research as goal. Organization, research and teaching. In 1990, the medical school (Faculty) and the hospital of the

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University of Amsterdam were the rst to form a University Medical Center: the Academic Medical Center of the University of Amsterdam, the AMC. By the late 1990s and early 2000s all Dutch Universities had followed. The basic sciences like physiology and anatomy originally departments of the University, became part of the Academic Medical Center or the University Medical Center, as well as the clinical departments. The research in physiology in the Netherlands is thus forced to make its research efforts more translational, and physiology as a stand alone scientic approach will probably disappear. I see a future for physiological research as a strong translational eld between genetic/ cellular research and its clinical applications, but not necessarily taking place in separate departments of physiology. The teaching in the University Medical Centres is more and more organized on the basis of organ systems, where cell, anatomy, physiology and clinical specialties all participate. This construction makes physiology as an entity less clear. However, the subjects should not disappear and be given by physiologists. Thus, the departments of physiology also have an opportunity to not only integrate their classroom teaching but also build up, together with the clinical departments, practical courses. A report released by the Association of American Medical Colleges (AAMC) in June 2009, entitled: Scientic Foundations for Future Physicians, emphasizes that physicians must have a rm grounding in the biomedical sciences and understand their relation to the physical sciences and mathematics (Long & Alpern 2009). This is an approach, where the teaching by the physiologist is essential. The physiologist as teacher should integrate these subjects to improve understanding the function of the organs and organism. The physiologists should also keep in mind that their teaching may help in the education of physicianscientists. Most often, physician-scientists either become pure clinicians or focus all of their energy on basic sciences, thus playing the same role as PhD-trained biomedical scientists, while they should form the core of the translational research effort. I urge the physiologists in the Netherlands to make a teaching programme that convinces the directorates of the University Medical Centers in the Netherlands that physiology is indispensable in the medical education. The teachings should not only confer essential physiological knowledge to all medical students but should also prepare, even a few of them, for future collaborative research between clinicians and physiologists. Departments may disappear but physiology does not and physiologists are indispensable.

Conict of interest
None.
The invaluable assistance of Connie Pieksma is greatly appreciated. I thank Professors Lennart Bouman, Gerrit Mook, Carel van Os, Dirk Ypey, and Marc Vos for their corrections and contributions.

References
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Acta Physiol 2011 Walter, W.G. 1961. Neergang van de physiologie in Nederland. Wolters, Groningen, The Netherlands. Westerhof, N. (2010). Hartog Jacob Hamburger Grand Dutch Physiologist. [WWW document]. http://www.feps.org/c Then click on Famous European Physiologists; then click Hamburger. (Last accessed 19 April 2011). N. Westerhof, Ph.D. Department of Physiology, VU University Medical Center, van der Boechorststraat 7, 1081 BT Amsterdam, the Netherlands. n.westerhof@vumc.nl

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