August and Marie Krogh August and Marie Krogh

LIVES IN SCIENCE

Bodil Schmidt-Nielsen, Dr. Odont, Dr. phil. Professor Emeritus and Aqjunct Professor, Department of Physiology, University of Florida

SPRINGER NEW YORK 1995 Oxford University Press

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Library of Congress Cataloging-in-Publication Data Schmidt-Nielsen, Bodil. August and Marie Krogh : lives in science by Bodil Schmidt-Nielsen. p. cm. Includes index. ISBN 978-1-4614-7530-9 (eBook) DOI 10.1007/978-1-4614-7530-9 1. Krogh, August, 1874-1949. 2. Krogh, Marie, 1874-1943. 3. Physiologists-Denmark-Biography. I. Title. QP26.K76S35 1995 591.1'092-dc20 [B] 94-20655

9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper Preface

When my father August Krogh died in 1949, 1 was with him in Den• mark. My stay in Denmark was prolonged for another two months due to a concussion 1 sustained in an automobile accident, which occurred shortly after his death. It was then that 1 became familiar with the ex• tensive material of biographical value which my father had left. Due to the pride and interest August took in his family, the material was very rich, consisting of family letters going back to his mother's birth in 1842, his own letters to his mother, father and siblings, between his siblings, between himself and my mother as well as his children. It in• cluded correspondence with his colleagues, university matters, Marie Krogh's letters, numerous notes and manuscripts, books, and scientific papers. It became clear that 1 was the one to write the biography because 1 was a physiologist. In addition, 1 spoke and wrote English as well as Danish and 1 very much wanted to undertake the project. My brother, Erik, was to inherit the material, but he was not inclined to undertake such a major job and he as well as Torkel Weis-Fogh (my father's scien• tific assistant) encouraged me to write the biography. It seems an awfully long time between making the decision and fin• ishing the work. 1 began immediately after having had the material shipped to Duke University, but soon found that it was impossible to combine such a major undertaking with my other work (research, teaching, and taking care of my three children). My resolve was undi• minished, but 1 had to put the writing on hold until my life became less complicated, which did not happen until after 1 retired. Weis-Fogh and 1 deposited the material in the Danish Royal Library in Copenhagen in 1964. During the intervening years, 1 continued to collect material of biographical significance. Then in 1987 the work began in earnest, but before 1 could start writing 1 had to copy all of the material we had deposited in the Danish Royal Library. Writing the book has been an absorbing and deeply rewarding un• dertaking. In studying my father's and mother's writings and correspon• dence, 1 have come to admire them even more than 1 had before. As my mind filled with what 1 read and then wrote, my parents carne vividly alive at all the various stages of their enormously productive lives. 1 have been asked if 1 learned things about them that 1 did not know before. 1 vi Preface certainly did, but the image of these two people who were my parents did not change, only the depth of understanding. 1 also arrived at the sobering realization that many thoughts, beliefs, and ideas that 1 had believed were original to me have come to me through what my parents taught me when 1 was a child.

Gainesville, Fla. B. S.-N. ]uly 1994 Acknowledgments

In this- undertaking I have had a great deal of help and encouragement. In copying the files from the Royal Library in Copenhagen, my grand• niece Maria Krogh Nielsen assisted me and Librarian Troels Lauring of Nordisk Insulin Laboratory kindly completed the task. All of it is now in my home in Gainesville, Florida. It is said that nothing is written until someone has read it. The first one to read my draft, encourage me and make small corrections was always my patient and beloved husband, Roger Chagnon. Next carne my excellent editor Miss Susan Williams who helped get the Danish out of my English, then carne my "two pro• fessors," Dr. Thomas Maren and Dr. Arthur Otis. I had told Tom Maren that I needed a critica! audience. He suggested forming a small commit• tee consisting of himself and Dr. Otis. During the years that I have been writing I regularly brought my pages to them. We would have lunch together while they read the next installment of the Krogh saga. I can• not thank these two friends enough for their never failing encourage• ment and kind suggestions. I have also regularly sent my manuscript to my family in Denmark, who have helped me with suggestions and more material and to Dr. Erik Hohwii Christensen in Sweden. Dr. Chris• tensen's review of my chapter on exercise was very helpful. Dr. Michael Bliss kindly reviewed the chapter on insulin and Dr. Hans U ssing the isotope chapter. My former teacher and lifelong friend, Else Rahbek, was always willing to help me contact people in Denmark or find mate• rial I needed. When I had almost finished, Dr. John West recommended the manuscript to the American Physiology History Section and to Oxford University Press, for which I am grateful. The last and least-pleasant part has been to condense my manu• script. I was told by the publisher it had to be shortened by about a third but given no specific guidelines for what to take out. An excellent editor in New York, Ms. Beverly Pennachini, has been sufficiently tough-minded to help me take out the less essential material. Contents

Chronology xi

Introduction 3

1. Ancestry 5 2. Childhood in Grenaa (1874-1890) 10 3. School Years in Aarhus (1891-1893) 17 4. Becoming a Zoologist (1893-1895) 22 5. Becoming a Physiologist (1895-1899) 32 6. The Beginning of Zoophysiology (1899-1904) 44 7. August and Marie (1904) 60 8. Marriage and the Seegen Prize (1904-1906) 71 9. The Oxygen Secretion Controversy (1906-1910) 78 10. The Zoophysiological Laboratory and the Beginning of Exercise Physiology (1910-1916) 95 11. Capillaries and the Nobel Prize (1916-1922) 112 12. The Insulin Story (1922-1925) 126 13. The Happiest Years (1923-1938) 139 14. Capillary Function and the Contractility Controversy (1923-1936) 154 15. New Fields of Investigation (1928-1943) 163 16. Exercise Physiology (1918-1944) 173 17. The Introduction of Isotopes (1935-1944) 185 18. Dark Clouds (1938-1944) 193 19. Exile in Sweden (1944-1945) 209 20. The Private Laboratory in Gentofte (1945-1947) 218 21. Last Scientific Work (1945-1949) 229

Epilogue 239 Notes and References 241 August Krogh's Publications 261 Index 277

Mimi Mimi

sisters sisters

1 1

brother brother

another another

7-1984) 7-1984)

4 4

Jens Jens

branch branch

1 1

Bodil Bodil

(Misse) (Misse)

Skovgaard Skovgaard

(194 (194

daughters) daughters)

(1784-1854) (1784-1854)

o) o)

(1915-) (1915-)

Emil Emil

2 2

Schmidt-Nielsen Schmidt-Nielsen

Marie Marie

Fischer Fischer

(1885-1952) (1885-1952)

Knut Knut

Jacob Jacob

1849-1933) 1849-1933)

( (

Ţ Ţ

Inger Inger

Brick Brick

Krogh Krogh

Freda Freda

(1821-191 (1821-191 1 1

(married, (married,

Christiane Christiane

(1944-) (1944-)

Mimi Mimi

Bent Bent

Marie Marie

(1788-1847) (1788-1847)

(1882-1922) (1882-1922)

(1918-) (1918-)

frida frida

Emil Emil

Ţ Ţ

-) -)

Bodil Bodil

1955) 1955)

1 1

Wernstedt Wernstedt

Dreckmann Dreckmann

Astrid Astrid

(1941 (1941

Jacob Jacob

1 1

(1822-1902) (1822-1902)

(?-1946) (?-1946)

1---

(1880-

Ditlev Ditlev

branch branch

Brewer Brewer

Christer Christer

(1945-) (1945-)

Andreas Andreas

Johan Johan

Christer Christer

daughter) daughter)

.------~-----.------, .------~-----.------,

Ţ Ţ

Tree Tree

1 1

Viggo Viggo

Baes Baes

de de

Helga Helga

(1848-1925) (1848-1925)

Brick Brick

son, son,

1 1

(married) (married)

(1 (1

Lars Lars

Andreas Andreas

Mygind Mygind

(1917-) (1917-)

(1939-) (1939-)

(1743-?) (1743-?)

Larsen Larsen

Krogh Krogh

Korsor Korsor

Maren Maren Bolette Bolette

branch branch

Agnes Agnes

Family Family

-Î-

Niels Niels

Ţ Ţ

(1778-1857) (1778-1857) (1781-1858)

Sofie Sofie

Detlef Detlef

(1918-) (1918-)

Christiene Christiene

Ţ Ţ

1 1

Bo Bo

Lauritz Lauritz

Krogh Krogh

(1878-1950) (1878-1950)

Hans Hans

Edel Edel

Krogh Krogh

Emil Emil

Johan Johan

Ljunghusen Ljunghusen

Andersen Andersen

(1816-1881) (1816-1881)

1 1

Andreas Andreas

(Line)-

Einar Einar

Jacob Jacob

Bjorn Bjorn

(1876-1950) (1876-1950)

(1942-) (1942-) (1948-)

Ţ Ţ

1 1

Christiansen Christiansen

Jacob Jacob

(1720-1786) (1720-1786)

Dreckmann Dreckmann

August August

Jorgensen Jorgensen

(1814-1867) (1814-1867)

1 1

Rigmor Rigmor

Marenline Marenline

Bodil Bodil

Christen Christen

(Mimi) (Mimi)

(1939-) (1939-)

(1913-) (1913-)

Marie Marie

(1874-1943) (1874-1943)

Ellen Ellen

Peter Peter

Dreckmann Dreckmann

branch branch

Birte Birte

Haders1ev Haders1ev

(1773-1846) (1773-1846) (1775-1815)

Bo1ette Bo1ette

(1799-1843) (1799-1843)

Hjort Hjort

1 1

Johann Johann

Ellen Ellen

(1944-) (1944-)

~ ~ (1914-) (1914-)

Karen Karen

SteenbergŢ SteenbergŢ

Ţ Ţ

Magdalene Magdalene

(1842-1927) (1842-1927)

.------,-----'-,------,-----,------, .------,-----'-,------,-----,------,

(Sine) (Sine)

Christensen Christensen

branch branch

Marie Marie

1 1

Viggo Viggo

August August

(1874-1949) (1874-1949)

(1770-1837) (1770-1837)

(1940-) (1940-)

Braenore-Frorup Braenore-Frorup

(1809-1876) (1809-1876)

Erik Erik

Inger Inger

(1908-1958) (1908-1958)

Christsigne Christsigne

Christian Christian Schack Schack Chronology

1874 August Krogh horn November 15 in Grenaa, Denmark. 1889-90 Voluntary apprentice of Danish Navy on the inspection boat Hauch. 1893 Artium (Entrance examination for the university) from Aarhus Cathedral school 1895 Preparatory examination for medicine, University of Copenhagen. 1899 Magister conference, University of Copenhagen. Scientific assistance in Christian Bohr's laboratory.

1902 First trip to Greenland to study atmospheric C02• 1903 Doctoral degree, Dr. phil. 1905 Marriage to Marie J0rgensen (horn December 25, 1874) 1906 Seegen Prize. 1907 Marie gets her medical degree. 1908 August and Marie study in Greenland. Erik Viggo Krogh is horn. 1910 Establishment of Zoophysiological Laboratory in Ny Vestergade. Publicq.tion of "the seven small devils." 1913 Ellen Rigmor Krogh is horn. 1914 Marie's doctoral degree, Dr. med. 1916 Professor of zoophysiology. 1917 Agnes Helga Krogh is horn. 1918 Bodil Mimi Krogh is horn. 1920 Nobel Prize physiology or medicine. 1921 Honorary Dr. jur., University of Edinburgh. 1922 Silliman Lectures at Yale University. 1923 Starts insulin production in Denmark. 1 1928 Establishment of the Rockefeller Institute in Copenhagen. 1935 Honorary Dr. med., University of Budapest. 1936 Honorary Dr. se., Harvard University. Honorary Dr. med., University of Lund. 1938 Honorary Dr. se., Rutgers University. 1943 Marie dies March 25. xii Chronology 1944-45 Exile in Sweden. 1945 Establishment of private laboratory in Gentofte. Baly Medal. Croonian Lecture. 1946 Honorary Dr. phil., University of Oslo. 1949 Resignation from the Danish Royal Academy of Sciences. Dies September 13, in his house in Gentofte. August and Marie 1\rogh