138 Lymphology 33 (2000) 138-140 BOOK REVIEW surprisingly little has been written about Starling's life, and equally remarkable he never received the Nobel Prize or was knighted unlike some of his British colleagues and students. Stimulated by these omissions, J ens H. Henriksen, a renowned clinical physiologist at Hvidore Hospital and the University of Copenhagen, decided on the centennial anniversary of Starling's now classical investigations into lymph formation to write a "short biography" of this investigator's extraordinary career. Raised in a family of 6 children with his British lawyer-diplomat father out of the country most of the time, Starling received a classical education including the German language. He initially qualified as a physician Ernest Henry Starling from Guy's Hospital but his major interest (1866-1927) was in physiology. He soon joined forces with by J ens H. Henriksen his brother-in-law, William Bayliss, to forge a Laegeforeningens Forlag professional collaboration which eventually Copenhagen, Denmark 2000 led to global acclaim for both men. Early on, Starling studied with Rudolf Heidenhain in Breslau, and it was that celebrated investi­ Whereas Assellius, Rudbeck, Pecquet, gator's misguided ideas that eventually led to Bartholin, Mascagni, William and John Starling's epic discourses on the origin of Hunter and several others are usually lymph while he was working at the University credited with the basic discoveries of the College in London (UCL). Although, as lymphatic system, it was Ernest Starling mentioned, Starling later discovered hormo­ through a series of epochal experiments in the nal secretions and the fundamental principle late 19th century who set the stage for the regulating the strength of myocardial modern understanding of the origin of lymph contraction, it was the "filtration hypothesis" and the key microcirculatory forces that is most meaningful to lymphologists. regulating the partition of the extracellular Beginning in the early 1890's, Starling fluid between plasma and the interstitial began his investigations into the dynamics of space. With Bayliss, Starling also introduced microvascular filtration and lymph forma­ the concept of hormones with the discovery of tion. Ludwig of Leipzig had earlier proposed secretin and promulgated the "law of the that lymph derived from plasma by filtration heart" regulating cardiac stroke volume. governed by arterial pressure. Two major Despite these monumental contributions, stumbling blocks, however, seemed incompa- Permission granted for single print for individual use. Reproduction not permitted without permission of Journal LYMPHOLOGY . 139 tible with that hypothesis. Thus, when composition in proteids of the transudate arterial pressure was artificially raised by fluid will vary ... we can therefore arrange the intravenous crude pituitrin (vasopressin), capillaries of the body in a descending order arterial hypertension followed but thoracic of permeability, the liver capillaries being the duct lymph flow either was unchanged or most permeable and the limb capillaries the decreased. Even more disconcerting was that least permeable" (2). when the inferior vena cava above the Henriksen succinctly relates these diaphragm or the portal vein was constricted, brilliantly conceived and imaginatively arterial hypotension was profound (as venous interpreted experiments to the understanding return and cardiac output decreased) but of edemas and coelomic effusions while thoracic duct lymph flow sharply increased. describing the resistance of many renowned Such discrepancies propelled Heidenhain into physiologists to explanations that seemed too proposing that lymphagogues (Le., vital mechanical and too simple. A list of forces) beyond physicochemical forces were Starling's most important and famous responsible for lymph formation. But Starling lectures on these subjects is also provided ingeniously reinterpreted the dynamic including the Arris and Gale Lectures (1984, responses of thoracic duct lymph flow 1896, 1897) and the Croonian Lectures in including its changing protein composition to 1904 and 1905. Although it took another 30 conclude that capillary pressure (in this years for the detailed microcirculatory instance selectively heightened either in the experiments by Pappenheimer, Soto-Rivera liver or extrahepatic portal bed), not arterial and Landis to verify the filtration hypothesis pressure, was the underlying mechanical almost in its entirety, Starling certainly force regulating blood capillary filtration and appreciated the clinical implications in lymph formation. Starling further realized almost every detail. As Cecil Drinker from these same experiments the importance expressed it (3), "those who have reworked of colloid osmotic pressure as a countervailing the field, though able to describe many things microvascular force-"although the osmotic which would have been immensely interesting pressure of the proteids of the plasma is so and gratifying to Starling, have added insignificant, it is of an order of magnitude comparatively little to the fundamental comparable to that of the capillary pressures; principles which he formulated of lymph and whereas capillary pressure determines formation and the movement of fluid from transudation, the osmotic pressure of the and into blood capillaries. Starling's evidence proteid of serum determines absorption... so though although technically crude, was that, at any given time, there must be a nevertheless the product of an intelligence balance between the hydrostatic pressure of which had a habit of being right in the great the blood in the capillaries and the osmotic essentials." attraction of the blood for the surrounding Using original diagrams to illustrate key fluids. With increased capillary pressure findings and innovative equipment utilized in there must be increased transudation, until these animal experiments as well as seldom equilibrium is established at a somewhat seen photographs of the laboratory and of higher point, where there is a more dilute Starling as a young investigator, Henriksen fluid in the tissue-spaces and therefore a captures the excitement and significance of higher absorption force to balance increased those halcyon days for clinical physiology and capillary pressure" (1). medical practice. Starling also appreciated the varied Several chapters are devoted to the permeability of the capillary membrane in discovery of secretin and investigations into different organs-"According to the perme­ the "law of the heart," which are also ability of the membrane, the amount and fascinating to read not only for the originality Permission granted for single print for individual use. Reproduction not permitted without permission of Journal LYMPHOLOGY . 140 of the experimental preparations but also for including a handwritten reproduction of the inspiration galvanized for future investi­ correspondence with August Krogh and a gations. Ironically, Starling's law of the heart compendium of official and unofficial is now thought to also govern propulsion of Starling memorial lectures delivered between lymph. Thus, distention of the Iymphangion 1963-1999. Starling's final years were stimulates a more potent thrust in lymphatic characterized by travel, declining health (? contraction and hence greater lymph colon cancer), and his last journey to the transport (the law of the "cor Iymphaticum"). West Indies where he died aboard ship and Other chapters dwell on Starling's his burial site in Kingston, 1 amaica. experience at Guy's Hospital, the constructive For the author Henriksen, this book was atmosphere for inquisitive learning, his time clearly a labor of love, packed with pictures, as 10drell Professor at UCL, the struggles vignettes, and even a color photograph of with antivivesectionists buttressed by cartoons himself on the back cover sitting in Starling's of the day and old and new photographs (the original chair at UCL. All in all, Henriksen latter taken by Henriksen) including has done an incalculable service to the Starling's office chair and laboratory (now a memory and contributions of Starling to conference room and library). clinical physiology and to Iymphology by Later chapters deal with Starling's putting together this historical tribute to one frustrations as a medical officer in World of England's and the world's greatest War 1. Despite his misgivings as an old (48 physician-scientists. As Lovatt Evans years) physiologist, Starling took a refresher eulogized Starling (4), "generalizations on the course in medicine and went to work on grand scale ... as opposed to niggling details strategic research to counteract the effect of always had a great fascination for him and it poisonous gas. Like the renowned physician was the noble and sweeping gesture of 19th and contemporary William Osler, Starling century biology [Darwinism] ... which expressed open hostility and disappointment impressed him more than the minutiae of at German aggression. Both men had had contemporary biological teaching." enormous respect for the German Universities REFERENCES and their progressive teaching policies and felt betrayed by the Kaiser and "Prussian 1. Starling, EH: On the absorption of fluids from power." Unlike Osler who lost his only son in the connective tissue spaces. J. Physiol. 19 (1896),312-326. that war, Starling had no children. Unfortu­ 2. Starling, EH: The Arris and Gale Lectures. nately, Starling found military politics The Royal College of Surgeons, 1894, 1896. overbearing,