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Bill Klug

Quiet Adventurer, Transcendent Human


It is with heavy hearts that we report the terribly

premature and tragic death of

Bill Klug

, a noted

practitioner of computational mechanics whose

interests centered on biophysical phenomena in

contexts ranging from viruses to the human heart.

It has been said that “life well lived is long,” but

Bill Klug’s incredibly well-lived 39 years were

far too short by any measure, regardless of how

extraordinarily well lived they were.

Bill was a California boy; it was his home from his

childhood years through college, graduate school,

and where he was now raising his own beautiful

young family. As an undergraduate at Westmont

College in Santa Barbara, and as a master’s stu-

dent at UCLA, Bill had already established two of

the great loves of his life, his wife

Mary Elise


the use of mathematics as a way to understand and

engineer the world around him. As a graduate

student at the California Institute of Technology,

where he studied with

Michael Ortiz

, Bill began

his first forays into the power of computational

mechanics as a way of studying the living world.

Bill’s thesis work was inspired by stunningly beau-

tiful single-molecule experiments that measured

how force builds up inside of viruses as a result of

their packaged genomes. Bill took a clever con-

tinuum mechanics approach for representing the

structure of the packaged DNA permitting him to

compute its stored energy.

One of Bill’s most impressive attributes — and

the basis of how he lived his life in general — is

that he was a quiet adventurer. Upon landing a

job at UCLA as an assistant professor in mechani-

cal engineering, nothing would have been easier

than to settle into a productive and even exciting

career studying traditional mechanics problems.

Instead, both in his teaching and research, he set

out to redefine what it means to be a computa-

tional mechanician, setting his sights on a quan-

titative understanding of the mechanics of living

systems, from sub-cellular scales to the scale of

human organs such as the heart.

Bill’s research focused on several areas of central

importance to biophysics, where it was clear that

the principles of mechanics played a critical part

in the underlying biological phenomena. Inspired

by his thesis work, Bill established many excit-

ing collaborations with experimental groups that

examined the mechanical response of viral capsids

to mechanical stimuli, such as those imposed us-

ing the atomic force microscope (AFM). In these

experiments, an AFM tip is used to push on the

virus, and Bill and his many collaborators used

a combination of analytical thinking and finite-

element models to study the mechanical response

of the capsids, making detailed comparisons with

the results of experimental studies. More recently,

Bill pioneered new numerical methods for the

application of non-linear elasticity theory to study

problems such as the maturation of viral capsids.

This led to a better understanding of viral capsids

as macromolecular machines that cleverly manage

built-in elastic stress.

The mechanics of lipid bilayer membranes was

another area that enticed Bill. Here too—whether

in the context of optical tweezer experiments on

membrane tethers or the study of the ordered

arrays formed by membrane proteins—he showed

how the finite element method could be used as a

powerful window onto membrane mechanics. He

was also intrigued by networks of cytoskeletal fila-

ments and made important contributions to our

understanding of the mechanics of networks of

stiff protein filaments, with implications for how

cells generate and respond to force. One of Bill’s

recent growing enthusiasms was the modeling of

the human heart. What captured Bill’s imagina-

tion in this problem was the interaction between

the complex geometry of the heart, its mechanical

response, and how these different aspects of the

heart dictate cardiac electrophysiology.

As all of these examples show, Bill’s mind traveled

adventurously across biological phenomena and

the length scales at which they occur. Further, his

talent and knowledge in attacking hard problems

with elegant and transparent numerical methods

coupled with his enthusiastic

Bill Klug