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OBITUARY Bruce D. Sidell 20 March 1948 – 8 February 2011 - The Journal of Experimental Biology 214, 2453-2454

July 28, 2011


The discipline of comparative and evolutionary physiology

suffered the loss of one of its most creative scholars earlier this

year, with the all-too-young passing of Bruce Sidell. As all who

knew Bruce ñ– either personally or from reading his many excellent

publications ñ– realize, his science exemplified the ë‘curiositydriven

í’ approach to investigation that commonly leads to the most

novel and fascinating of discoveries. Bruce marched to the beat of

his own drum, and the pathway he followed throughout his career

was intellectually, technically and geographically wide-ranging and

adventurous. He spotted questions that others in his field had

missed and taught us many lessons about ë‘how animals workí’ that

we otherwise might not have learned. Bruceí’s approach to science

also taught us much about the importance of collegiality, candor

and collaboration in advancing the scientific enterprise. Bruceí’s

personal model not only helped to shape the career approaches of

his many graduate students but also served as an exemplar to his

peers and colleagues of how best to pursue a life in science. In

preparing this tribute, we were again reminded of all that Bruce

meant to us as a friend, collaborator and intellectual catalyst of his

field. We hope that our words serve as a clear reminder not only

of what Bruce discovered but also of what he taught us about the

most effective ñ– and enjoyable ñ– modes of fostering discovery.

Throughout his scientific career, Bruce manifested a remarkable

talent for formulating novel and important questions and then

choosing the right organisms and methodologies to get the answers.

The science from his laboratory was always well crafted in design,

executed with precision, and interpreted with the right balance of

conservatism and speculation. His success is reflected in a career

of uninterrupted funding from the National Science Foundation

(USA). The output from his laboratory at the University of Maine

was impressive in quantity and quality and continues to serve as an

intellectual springboard for studies on a wide spectrum of questions

in thermal biology.

Bruceí’s decision to pursue a career in biology can probably be

traced to his introduction to research at his undergraduate alma

mater, Boston University, in the laboratory of Frank Belamarich,

whom he admired greatly. From there, Bruce moved to the

laboratory of C. Ladd Prosser at the University of Illinois, which

at the time was a leading center for study of the thermal biology of

animals. Bruceí’s primary interest quickly became the impact of low

temperature on the biochemistry of fish. Whereas he cut his

intellectual teeth in work with temperate eurythermal species, most

of the latter half of his career was spent investigating Antarctic fish

that live close to the freezing point of seawater and die at

temperatures only a few degrees above zero. Bruceí’s curiosity led

him to investigate a great many facets of the thermal biology of

these and other species of fish. Indeed, much of Bruceí’s success

may be attributed to his ability to appreciate the full jigsaw puzzle

of thermal biology, from biochemical and molecular mechanisms

to physiological ecology and evolutionary biology. He was able to

work on sections of the puzzle throughout his career, often at the

same time, but most important of all, he saw how the pieces fit

together with an intellectual skill that elevated him above his peers.

The combination of exquisite experimental detail and over-arching

intellectual synthesis that one finds in Bruceí’s publications is

remarkable and truly the hallmark of his contributions to our field.

Bruceí’s first interest, and one that continued throughout his life,

was energy metabolism. Work in this area included his

contributions to metabolic fuel preference at low temperature, the

role of fatty acid binding proteins, and the control of lipid

degradation at the hormonal and enzyme levels. Just as examples,

his laboratory was the first to compare metabolic enzyme capacities

in laboratory-acclimated fish with those in field-captured fish, and

the first to show the metabolic preference for unsaturated fatty acids

in Antarctic fish. Recognition that ATP supply was only one

component of the metabolic story set the stage for experiments

related to energy demand in red and white muscle. His integrative

studies on striped bass, which covered isolated contractile proteins,

biophysics of contractile fibers, fiber recruitment at the wholeanimal

level, and calcium management provided the first

comprehensive picture of the impact of temperature on swimming

performance. A third broad area that captured Bruceí’s interest, one

in which he again filled in key pieces of the puzzle about

mechanisms of adaptation to low temperature, was how oxygen

moves from the sarcolemma to mitochondria. Conceptual

advancements on this front included ultrastructural modifications

at the mitochondrial level to reduce diffusion distances and the

exciting and novel idea that lipid pools may serve as a conduit for

oxygen movement. He demonstrated lipid-mediated oxygen

transport in an elegant series of experiments, using a device he

designed and fabricated himself.

Deeper understanding of oxygen delivery was enhanced with a

series of papers on myoglobin function that elegantly exploited the

The Journal of Experimental Biology 214, 2453-2454

© 2011. Published by The Company of Biologists Ltd



Bruce D. Sidell

20 March 1948 – 8 February 2011

Bruce Sidell on Cormorant island, Antarctic Peninsula, with Mt William in

the background, in 2009. Photo courtesy of Dr Jody Beers.

􀀴􀀨􀀥 􀀪􀀯􀀵􀀲􀀮􀀡􀀬 􀀯􀀦 􀀥􀀸􀀰􀀥􀀲􀀩􀀭􀀥􀀮􀀴􀀡􀀬 􀀢􀀩􀀯􀀬􀀯􀀧􀀹


large variability of this protein among species. Bruce focused

strongly on the hearts of Antarctic icefish, some of which lack

myoglobin entirely. He elucidated the different mechanisms

underlying the loss of this putative ë‘essentialí’ component of the

oxygen transport system, showing that different molecular lesions

were found in different lineages. Using a combination of

physiological approaches, he showed that loss of cardiac

myoglobin reduced cardiac aerobic capacity. He also studied the

function of myoglobins in these highly cold-adapted fish,

demonstrating that these proteins had an unprecedented ability to

unload oxygen rapidly at extremely low temperatures.

Bruceí’s final line of investigation perhaps most clearly illustrated

his ability to ë‘connect the pieces of the puzzleí’ in an insightful and

synthetic fashion. These efforts were directed to building an

integrated perspective that comprises the roles of heme-containing

proteins (hemoglobin and myoglobin), the production of nitric

oxide, and angiogenesis. He built a strong case that the lack of

heme-containing proteins in icefish resulted in decreased

conversion of nitric oxide (NO) to nitrate, leading to higher steadystate

levels of NO that subsequently triggered both mitochondrial

proliferation and angiogenesis. In this way, he showed the

evolutionarily downstream consequences of the loss of hemoglobin

and myoglobin in this lineage of Antarctic fish. His later studies

also revealed the importance of oxygen-carrying pigments in

setting the upper thermal tolerance limits of Antarctic fish. He

showed that the hemoglobin-less icefish are much less tolerant of

elevated temperatures than their red-blooded relatives.

Consequently, icefish may be especially susceptible to the effects

of climate change.

It was this type of multi-level, highly integrative thinking that

put Bruce a step ahead of his peers. Although Bruceí’s experiments

were often specifically focused on questions in the context of cold

thermal biology, the conceptual impact was far reaching in yielding

insights into fundamental principles, especially as related to

mechanisms of metabolic control, muscle performance, and oxygen

delivery. His contributions thus taught us a great many new things

about nature and, equally importantly, illustrated the types of

approaches to research that are most fruitful and fun. We also note

that he was as comfortable on a freezing deck trawling for

specimens off the Antarctic Peninsula or the South Shetland Islands

as he was conducting experiments in his laboratory in Maine or at

the U.S. Antarctic base at Palmer Station.

It would be unfair, and certainly incomplete, to restrict our

comments to the strictly scientific side of Bruceí’s contributions. He

was an academic man in full, a strong contributor to the teaching

and administrative activities of his university and an important

participant in the Antarctic scientific community, including his long

service to the Palmer Area Users Committee and the Antarctic

Research Vessel Oversight Committee. In the latter role, as the sole

􀀁 scientific advisor on a $100million future facilities contract, he had

to maintain his characteristically balanced and objective approach

in the face of political pressures from industry and legislative

constituencies. In recognition of his long career of research and

service to the Antarctic scientific community, in 2010 the Sidell

Spur on Brabant Island in the Palmer Archipelago, overlooking an

area where he often fished for specimens, was named for him by

the National Science Foundation and the United States Geological


Bruceí’s entire career as a professor was spent at the University

of Maine. There, he taught a series of highly acclaimed

undergraduate (Cell Biology; Vertebrate Biology; Biology of

Fishes; Integrative Marine Science) and graduate (Physiology of

Fishes; Biochemical Adaptation) courses and mentored generations

of graduate students, who are now successful in their own rights.

He did yeoman service as the founding Director of the School of

Marine Sciences, where he brought together a diversity of marine

biologists, oceanographers and aquaculturists, as well as

economists and policy and management specialists, to form an

academic department that is consistently at or near the top in

research funding and productivity at his university. Bruceí’s success

in administrative tasks was based on the same personal

characteristics that helped account for his success as a scientist:

high intelligence, extraordinary attention to detail, an ability to

inspire loyalty and a high level of effort in the colleagues and

collaborators he chose to work with, and a unique sense of humor

that helped him bear the diverse burdens associated with academia.

In recognition of the esteem in which he was held, his friends,

colleagues, and former students established the Bruce D. Sidell

Scholarship for deserving students in the School of Marine


Bruceí’s success as an academic was paired with a rich and

supportive family life. He enjoyed the companionship of a

wonderful wife, Mary, their three daughters, Amy, Jessica and

Amanda, and their husbands, and, later, six grandchildren. Bruce

would also want us to recognize his canine companions who, along

with his family, carried him through the challenges of his

professional life.

Bruce Sidell manifested the mixture of deep curiosity about

nature, creativity in experimentation, cooperativeness in working

with colleagues, caring and fairness in mentoring students, and

commitment to the diverse expectations of academic life that

underlies a most successful career. Proud of his liberal education,

Bruce was mindful of his broader audience and the role of science

in society, and had underlined these words in his copy of William

Zinsserí’s book ë‘On Writing Wellí’: ë‘Ö…they all come across first as

people: men and women finding a common thread of humanity

between themselves and their specialty and their readers. You can

achieve the same rapport, whatever your subject.í’ His passing will

be deeply felt by all in our field, but his rich legacy will continue

to shape our thinking about our science and inspire us to carry out

our activities in research and education in the creative and

thoughtful way that Bruce exemplified.

William Driedzic

Ocean Sciences Centre, Memorial University of Newfoundland

J. Malcolm Shick

School of Marine Sciences, University of Maine

George N. Somero*

Hopkins Marine Station, Stanford University

*Author for correspondence (


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