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101

SYMPOSIUM
Bengt Saltin and exercise physiology: a perspective1
Michael J. Joyner

Abstract: This perspective highlights some of the key contributions of Professor Bengt Saltin (1935–2014) to exercise physiology.
The emergence of exercise physiology from work physiology as his career began is discussed as are his contributions in a number
of areas. Saltin’s open and question-based style of leadership is a model for the future of our field.

Key words: work physiology, thermoregulation, muscle glycogen, exercise capacity, muscle plasticity.

Résumé : Cet article souligne quelques-unes des contributions essentielles du Pr Bengt Saltin (1935–2014) à la physiologie de
l’exercice physique. On analyse l’émergence de la physiologie de l’exercice issue de la physiologie du travail au moment où sa
carrière commence en plus de ses contributions dans plusieurs domaines. Son leadership axé sur l’ouverture et le questionne-
ment constitue un modèle pour l’avenir de notre domaine. [Traduit par la Rédaction]

Mots-clés : physiologie du travail, thermorégulation, glycogène musculaire, capacité d’effort, plasticité musculaire.

Introduction the early focus on economic as well as biomedical issues. In this


This perspective is about the impact and career of the Scandina- context, an outstanding example of classical work physiology is a
vian physiologist Bengt Saltin (1935–2014). It is based on a presenta- study on thermoregulation and adaptation to heat stress con-
tion given at the 2015 Canadian Society of Exercise Physiology ducted by Sid Robinson at the Harvard Fatigue Lab during World
meeting that was part of a symposium that included lectures on War II. At that time there was concern that soldiers from cool
Professor Saltin’s larger scientific contributions and his interac- climates would not be able to deal with the heat in tropical envi-
tions with Canadian scientists. That having been said I want to ronments. This study showed that several weeks of daily exercise
frame the perspective via 3 main questions: First, what was the in a warm environment evoked marked improvements in thermo-
exercise physiology world like and where did it come from when regulation and blunted some of the concerns about tropical envi-
Bengt Saltin started in research in the early 1960s? Second, what ronments (Robinson et al. 1943).
do the highlights of his career tell us about our field and what The central role of work physiology in our field is further high-
lessons do they offer today? Third, what can we learn about the lighted by the title of one of the foundational texts in our field, the
leadership of scientific enterprises from reviewing Bengt Saltin’s Textbook of Work Physiology (Åstrand and Rodahl 1970) was first
career? published in 1970 and coauthored by one of Bengt Saltin’s teach-
ers and early collaborators Per Olaf Astrand. It should also be
Exercise Physiology before 1960 noted that the pre-World War II “Work Physiologists” were cer-
When I talk to students about where exercise physiology came tainly interested in exercise and incredibly insightful studies on
from, I always emphasize our links to what might be described as topics, such as maximal oxygen uptake, exercise metabolism, and
industrial or work physiology. Starting in the late 1800s and well human performance were well underway during that era (Robinson
into the 1900s, there was great interest in these topics because et al. 1937). These studies also set the stage for what followed espe-
many humans in the “developed” world worked at jobs that re- cially in the 1950s, 1960s, and 1970s. It is important to note that up to
quired extensive manual labor. As the world industrialized there that time a lot of attention had been paid to the role of the heart and
was an economic and political need to understand the energy cardiac function as a key determinant of exercise capacity.
requirements of this work, how it was fueled, and how it was
affected by environmental conditions. At the same time there was Lessons from Saltin’s contributions
a larger hygiene movement focused on improving living and I like to say that I first met Bengt Saltin in the late 1970s. How-
working conditions for the masses. Additionally, heroic feats of ever, that meeting was really a meeting of his ideas and occurred
physical endurance, such as polar exploration and mountaineer- via reading and discussing his papers with fellow students in a
ing, captured the imagination of the general public as mass com- journal club at the University of Arizona that was led by Jack
munication emerged. Wilmore. Three of the major themes of his work were already well
The names of 2 of the earliest North American programs on developed by then and they include thermoregulation, maximal
topics related to our field reflect these trends. At the University of oxygen uptake, and regulation of skeletal muscle metabolism.
Toronto there was a department of Physiological Hygiene and the One of his earliest papers on the impact of dehydration on maxi-
Harvard Fatigue Laboratory was founded in Boston in the 1920s mal oxygen from 1964 was discussed (Saltin 1964). The paper
(Best 1928; Scheffler 2015). That the Harvard Fatigue Lab was lo- shows that dehydration had a much bigger impact on time to
cated in the business school and not the medical school highlights exhaustion than maximal oxygen uptake. It remains a classic and

Received 6 June 2016. Accepted 1 September 2016.


M.J. Joyner. Department of Anesthesiology, Mayo Clinic, Rochester, MN 55905, USA.
Email for correspondence: joyner.michael@mayo.edu.
1This paper is a part of a symposium entitled A Tribute to Bengt Saltin: The use of Exercise to Study Integrative Physiology.

Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink.

Appl. Physiol. Nutr. Metab. 42: 101–103 (2017) dx.doi.org/10.1139/apnm-2016-0314 Published at www.nrcresearchpress.com/apnm on 12 September 2016.
102 Appl. Physiol. Nutr. Metab. Vol. 42, 2017

has all of the hallmarks of his later work, including a clear hy- Fig. 1. Key topics covered in Saltin’s inaugural Wolffe Lecture.
pothesis, straight forward experimental design, and a highly in-
formative and succinct discussion. Typically, figures that were
simple but elegant reinforced the key findings and interpretation.
By the late 1960s Bengt had begun to focus on muscle metabo-
lism and he was one of the authors along with Bergstrom, Her-
mansen, and Hultman on the effects of diet on muscle glycogen
and performance. This study leveraged the muscle biopsy tech-
nique and showed that muscle glycogen stores were a key deter-
minant of endurance time during cycling at about 75% of maximal
and set the stage for practices like carbohydrate loading and feed-
ing during exercise (Bergström et al. 1967). In addition to spawn-
ing a vast array of science, this study (and others) was critical in
sparking the whole field of sports nutrition and the downstream
commercial efforts to improve performance via various drinks,
gels, and supplements.
In 1972 Bengt presented the first Wolffe lecture at the American
College of Sports Medicine. The title of the lecture and subsequent
paper published in 1973 was Metabolic Fundamentals in Exercise
(Saltin 1973).
Figure 1 is simply a listing of the subheadings of that paper. I
include it to show the breadth of Bengt’s thinking and to high-
light that many of the topics he discussed remain vibrant areas of history and scientific legacy of the CMRC has been reviewed in
ongoing research in our field. The paper is also worth rereading detail elsewhere (Joyner et al. 2015). However, as someone who
from time to time as a reminder of what we do and do not know was not a member of the CMRC and never worked there several
and as a reminder of the timeless nature of good questions. Here things stand out. First, the CMRC was highly collaborative. On a
is a quote from the paper that I particularly like: scientific basis this permitted collaborations in areas at the inter-
It is tempting to suggest that the relative muscle fiber com- section of topics such as blood flow and metabolism to thrive. This
position be used as a method for identifying early in life collaborative spirit also attracted top young investigators from all
those able to compete successfully in different physical ac- over the world who came for training and collaborations. Second,
tivities or to predict the potential to adapt to training that activities at the CMRC also led directly or indirectly to the gener-
the muscles and body may have. Such identification in no ation of a number of outstanding international symposia and
way, however, obviates the need for intense training. courses. It was through these events that I got to know Bengt
This quote anticipates by about 40 years the current focus on the Saltin personally and interact with him. Bengt was unfailing in his
molecular and other omic factors that along with concepts from inclusion and encouragement of young investigators from all over
traditional integrative physiology explain inter-individual differ- the world. I also saw firsthand a leadership style that was open,
ences in the responses to exercise training. It also anticipates the question-focused, and flexible versus excessively hierarchical and
rise in things like direct to consumer testing for talent identifica- bureaucratic. At some level the best labs are like Montessori
tion (Webborn et al. 2015). schools for bright young people. Bengt showed it was possible to
The next big wave of innovative ideas from Bengt Saltin’s fertile run a large and high-powered research center at a major research
mind came in the 1980s with the development in Copenhagen of university in the same way.
an improved thermodilution technique to measure leg blood flow
in combinations with “1-leg” kicking exercise. These techniques Summary
essentially permitted the quadriceps muscle to become a human The career of Bengt Saltin highlights the emergence of exercise
bioassay system. This technique has been essential in understand- physiology and sports science from its roots in work physiology.
ing the regulation of skeletal muscle blood flow. Most importantly His career also shows that it is possible to do mechanistic studies
data collected with it showed blood flow to human skeletal mus- in humans by combining a suite of techniques ranging from clas-
cles could be much higher than previously accepted values (250– sical measurement of gas exchange to molecular signaling. To do
400 mL·100 gm−1·min−1 vs. 50–100 mL·100 gm−1·min−1) (Andersen this requires a question versus technique centric focus. Bengt’s
and Saltin 1985). This observation was also central to raising and career is also a testament to the power of collaboration with oth-
addressing issues about the need for the autonomic nervous sys- ers and facilitation of their work. In physiology we frequently
tem to control the distribution of cardiac output and skeletal discuss feedback control. Bengt Saltin’s contributions, achieve-
muscle blood flow during whole-body exercise for the purposes of ments, and leadership represent 50 years of positive feedback for
regulating blood pressure. The blood flow technique was also the field of exercise physiology.
combined with the muscle biopsy technique and microdialysis so
that complex issues at the interface of blood flow and metabolism Conflict of interest statement
could be addressed. The advantages of this human model include Michael J. Joyner, MD, reports no conflicts of interested associ-
the ability of volunteers to cooperate while performing complex ated with this manuscript.
tasks, studies in patients, and also the simple fact that the quad-
riceps is a big muscle that facilitates sampling. These issues and References
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(Joyner and Casey 2015). 669 pp.
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