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BESSEL KOK
November 7, ~91~—April 27, ~979
BY J. MYERS
TH E D E C A D E ~ 955-l 965 was a period of revolution in
our thinking about the process of photosynthesis. A key
event was the realization that the process, long assumed to
require only one photoreaction, actually required two. A key
figure in the revolution was Bessel Kok.
In the following years Bessel and his collaborators in
work that bore the stamp of greatness Flee in much of the
framework for the Z-scheme mocle! of the energetics of pho-
tosynthesis. A preclictable result was the recognition that fol-
lowecI. There was a Kettering Research Award in 1963 given
by the Charles F. Kettering Foundation ancI the National
Academy of Sciences. There were two awards from the
American Society of Plant Physiologists: the Charles F. Ket-
tering Awarct in 1972 and the Stephen Hales Awarcl in 1978.
Anc! there was election to the National Academy of Sciences
in 1974.
In composing this memoir for Bessel Kok ~ shall use
freely the thoughts of collaborators, family, and friends,
which are recognizes] only partially in the acknowlecigment
at the end. ~ was for almost thirty years a distant colleague,
a sometime confidant, a scientific admirer, anct a friend all
of which accounts for the highly personal tone of this ac-
125
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126
BIOGRAPHICAL MEMOIRS
count. But it wouIcl be difficult for anyone to write otherwise
because Bessel was a very personal person.
Bessel Kok was born November 7, HIS, in the village of
Hardinxveld, The Netherlands. His father, Tohannes Evert
Kok, was the principal of a local school; he is remembered as
a talented man who cast himself in the image of a professor,
a puritan Calvinist who lived at the hand of the Bible. His
mother, Cornelia Gronclys-Kok, is remembered as "the image
of Bessel in female form." Bessel was the oiliest in a family of
six chilctren. He remembered his chilc~hood as happy, al-
though it was interspersed with the many frustrations that
must have arisen for an inventive and energetic boy encum-
berecl by a conservative father.
Bessel's own statement about his education was that he
"never stuctiect" and that he coasted through his high school
and college years. Nevertheless, he assimilated a broad back-
ground in science anct cleveloped the self-ctiscipline he
neeclec! thereafter. His college years began at the University
of Leiclen in 1934; they led to the unclergraduate degree of
Candidate of Natural Philosophy in 1938 and the advanced
degree of Doctor of Natural Philosophy in 1941. An even
more significant event occurred! in 1938 when Bessel met
Cornelia Hendrika Vogelesang at a Christian Stuclent's Club.
Cornelia or Nell, as she became known to all hac! been
born and raised in TancIjung Pinang, Indonesia. Her mem-
ory of meeting Bessel is one of instantly recognizing him as
her future husband.
In 1941 Bessel began work toward a Ph.D. in biophysics
at the University of Utrecht. In that era the road towarc! a
degree was tortuous and at times must have seemed impos-
sible. Following the German occupation in 1940, young men
were conscripted for labor camps in Germany. But for all it
was a time of struggling merely to survive. To provide some
protection against conscription, Nell and Bessel were mar-
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BESSEL KOK
127
ried in 1943. And although she was selclom successful, Nell
clid her best to hicle Bessel. He, on the other hanct, was more
inclined to trust to his luck anct the weak support of a pass-
port with a falsely increased statement of age. Bessel had
become assistant manager of a distilling company, Johan Kos-
ter. He also improvised a small ctistillery at home to produce
gin from rye and beets; the gin was readily bartered for food.
Their first chilcI, Lily, was born in March of 1945, two months
before the end of the war. Nell remembers it as the "worst
time" for an impoverished and starving people.
With the ens! of the German occupation, Bessel was free
to turn his efforts from survival to science. The firm of Than
Koster provided him both with employment and some direct
support of the research for his dissertation, which was pre-
sented early in 1948. In the foreword of the dissertation,
Bessel recognizes with thanks his professors Bungenberg do
Jong and Baas Becking at Leiden. Professor Koningsberger
is namect as the Hooggeachte Promotor, but it is also maple clear
that E. C. Wassink was the real supervisor. Hence, Bessel's
"scientific genealogy" traces to the Biophysical Research
Group uncler the direction of A. I. Kluyver and J. M. W.
Milatz.
Bessel's dissertation was a study of the quantum yield of
photosynthesis in the alga Chlorella. It is not now, and was not
then, a very exciting document. One clifficulty was that the
subject itself, which had been hot in the early 1940s, was only
smoldering in 1948. A second difficulty was that a quantum
yield, usually expressed as the reciprocal or quantum num-
ber (quanta absorbed per oxygen molecules evolved), is ac-
tually no more than a number. Its validity hinges only on the
nitty-gritty details of measurement.
It must be a source of amazement to much of the scientific
worict that the quantum yield! of photosynthesis should have
engendered so many man-hours of work and yet so much
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128
BIOGRAPHICAL MEMOIRS
controversy. Certainly one of the many aspects of an expla-
nation of this circumstance is historical. In 1923 Otto War-
burg tract reported a minimum quantum number of 4.3, nat-
urally interpreted as 4. In its time this was an heroic
accomplishment. Warburg had invented a manometric
method for the measurement of gas exchange. Anct he tract
introduced a convenient plant material, the alga Chiorella.
There were several acictitional features of experimental pro-
toco! that turned out to be important. The first was the use
of optically clense cell suspensions that absorber! virtually all
the light. Hence quanta absorbed from a monochromatic
light beam couIcl be counted simply as incident quanta. The
second feature was an assumption that oxygen evolution of
photosynthesis was properly evaluated from pressure
changes observed in short light periods minus those observed
in alternating short dark periods.
Considering the theoretical significance of the quantum
number, it is remarkable that Warburg's value went unchal-
lenged for some fifteen years. By the early 1940s, however,
other measurements hac! been made. The number 4 was in
doubt as being too low, and the special experimental protocol
was being questionecl.
Bessel's choice of experimental conditions shows his in-
sight anct understanding of the problem. First, he maintained
conditions of steady-state photosynthesis and measured ox-
ygen evolution over a long time period (an hour). Seconcl, to
ensure steacly-state conditions in all cells, he used optically
thin suspensions. (Otherwise, cells in a shaken suspension
wouIcl alternate between periods of light and virtual dark-
ness.) This required mastery of the technology of the Ul-
bricht sphere, a device for measuring fractional absorption
by a light-scattering cell suspension. A third important choice
was that for each cell preparation he measured rates of ox-
ygen evolution at several different light intensities. Then the
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BESSEL KOK
129
slope of the expected linear plot of oxygen per second versus
absorbed quanta per second would give the quantum yield.
The quantum numbers obtained from forty-two sets of mea-
surements fell within the range 6.5 to 10.0. Bessel deduced
that 7.5 was "the most favorable value."
In addition to fincting that the quantum number was high
(7 to 10) rather than low (4 to 5), Bessel discovered a related
phenomenon. The curve for the oxygen rate versus the
quantum rate hac! two linear segments with slopes in the ap-
proximate ratio of Al and converging near the compensa-
tion point where photosynthesis just balances respiration.
The phenomenon, which has come to be called the Kok ef-
fect, is commonly thought of as resulting from a "suppression
of respiration." It provides a possible explanation for low
quantum numbers (as 4 to 5) observer! at very low light in-
tensities below the compensation point.
In 1949, Bessel joined the Solar Energy Research Group
of the Organization for AppliecT Scientific Research (T.N.O.)
under E. C. Wassink at the Agricultural University in Wag-
eningen. This was an excellent match for his needs. As he
later wrote on an employment record, it provicled for "full
time research, freedom, adequate services." A major mission
was the mass culture of algae, a subject that Bessel followed,
albeit sporadically, for the rest of his life. An immediate ques-
tion was the maximum efficiency of Chlorella in producing
total cell material, the efficiency for growth. Bessel's result
was an efficiency of about 20 percent (equivalent to a quan-
tum number of about 10), which still stancis as a benchmark.
Simply growing algae, however, was too bland for Bessel's
taste. Most of his efforts went into attendant basic problems.
There is an ultimate limitation on the yielcl of algae that
can be achieved under sunlight illumination. The problem is
that photosynthesis and growth become rate limitect at a light
intensity far less than that of midday sunlight. Those of us
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30
BIOGRAPHICAL MEMOIRS
concerned with mass culture naturally sought ways to circum-
vent the limitation. Of several possibilities the most interest-
ing was to take advantage of the intermittent light effect of
photosynthesis. With sufficient turbulence in a dense culture,
individual algal cells move rapidly into and out of the illu-
minated front surface and thereby receive high light only in
flashes. It was well known that short flashes could be used
with higher efficiency, but the time parameters of the effect
were not known. Bessel set out to study photosynthesis in
flashing light.
Bessel spent much of 1951-1952 on leave from Wagen-
ingen and as a fellow of the Carnegie Institution in the De-
partment of Plant Biology at Stanford under Stacy French.
The family now with two chilctren, Lily and young Besse}-
livect in the oic! barracks-like buildings that later became the
Stanford Research Institute. ~ first met them there and was
attracted by the happy self-sufficiency of a family learning
the ways of a strange land. Bessel's part of the laboratory
became a shambles of equipment-building, and he was frus-
tratecT by the slow progress of a machinist who was construct-
ing a sector for light chopping. His work at Stanford was
reported in a chapter of the 1953 Carnegie monograph on
algal culture, and it provider! at least a partial answer: any
reasonable turbulence conic! be expected to give some gains
in the yield of an algal culture under sunlight. Large gains,
requiring very short flashes, probably would be impractical
because of the power cost for the necessary turbulence.
On his return from Stanford, Bessel began extending his
work with flashing light to an attack on the kinetics surround-
ing the photochemical events in photosynthesis. His remark-
able experimental talents now came into play. He used a high-
intensity projection beam that could be chopped and/or
attenuated there were two coaxial sectors that allowed in-
(lependent variation of both the light and the clerk periocls.
.
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BESSEL KOK
13
AS a side effort, he hac! cleveloped almost to ultimate lim-
its—the measurement of gas exchange by volumetry. Now he
couIct choose a very small reaction vessel (<5 mm diameter
coupled to a volumeter arrangement that measured oxygen
evolution with high sensitivity. The broacl scope of the ex-
periments proviclect an unequivocal answer to a twenty-year
controversy about the reality of the "photosynthetic unit."
The oxygen yield from short flashes confirmed the earlier
fincting of R. Emerson and W. A. ArnoIcI: a maximum flash
yielct inclependent of temperature and equivalent to about
one O2 per 2,000 chlorophylls. Lengthening the flashes gave
a temperature-dependent flash yield approaching the rate
observed in continuous saturating light. Bessel again went
beyond the immediate anc! obvious objective in order to ob-
tain an explanation of earlier and apparently conflicting re-
sults.
In 1955 Bessel again visitec! the United States for two
meetings: a Gatlinburg conference on photosynthesis fol-
lowocl by a woric! symposium on applied solar energy at Tuc-
son and Phoenix. E. C. Wassink also was in attendance. It was
evident to all, however, that Bessel tract attained the status of
an inclepenclent and innovative scientist.
The sectors of Bessel's flashing light apparatus must never
have coolecl (town for very long. Within a year after the mea-
surement of oxygen flash yielcis, the apparatus was recle-
signecI. Now the reaction vessel was a cuvette that was cross-
illuminated by a weak measuring beam from a monochro-
mator. Any small changes in the transmission of an algal or
chioroplast suspension couIcl be observed by a photocell.
Were there absorption changes that might reveal photochem-
ical intermediates formed cluring a short flash and removed
cluring the following ciark period? There were. Actually
there were several absorption changes observable across the
spectrum. Bessel zeroed in on one of these, a reversible ab-
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132
BIOGRAPHICAL MEMOIRS
sorption change at 700 nm that he speculated might be the
"eventual final photoreceptor of photosynthesis." At this
point the tribute of Lou Duysens is appropriate: "The long
working hours and inexhaustible inventivity and drive nec-
essary to get such an apparatus working successfully in a rel-
atively short time, anct to carry out ant] analyze a large num-
ber of experiments, were amply rewarcled....Besse}
ctiscoverect the far-red absorption changes associated with the
reaction center P700 of system IN
By 1957 Bessel's accomplishments had become highly vis-
ible. He had outgrown his position at Wageningen, and no
appropriate position in The NetherIancls was open. Like
many Dutch scientists before him, Bessel tract become avail-
able for export. A position as clirector of a new institute in
West Germany was offered; Bessel ancT Nell visited and to-
gether concluded that they shouIcT reject the offer. One major
consideration was that the rigidity and formal protocol of the
establishment was incompatible with their own life-style.
Among other inquiries was one from the Research Institute
for Acivancect Studies (RIAS) in Baltimore. By 1958 the Kok
family now with three children—hacI found a new home in
northwest Baltimore, not far from the converted mansion
that housed the RIMS laboratories.
RIAS was then about two years oIc} and not well known
even in the United States. It was a subsidiary of the Martin
Company, a corporate investment as an institute for basic
science. In the course of a European trip, its director, Wel-
come Bencler, tract stopped to see Bessel in Wageningen, anc!
his recruitment was that simple. Bessel electecT to accept a
position as a "staff scientist" in an unproven, industrially sup-
ported institute. There were no trappings to the position-
' L. N. M. Duysens, "In Memory of Bessel Kok," in Proceedings, Fifth International
Congress on Photosynthesis, ed. G. Akoyunoglou (Philadelphia: Balaban International
Science Services, 1981), pp. xix-xx.
OCR for page 133
BESSEL KOK
133
not even the conventional one of academic rank but only
the promise of logistic support and freedom of choice in re-
search.
Bessel tract joined a small group labelled "Bioscience" and
with a common bond of interest in photosynthesis. Within
the year he tract become the cle facto leacler of the group. No
one worked harder, sprouted ideas faster, or drank more
martinis at Friday afternoon parties. Welcome Bender man-
aged the whole institute with the light touch of one who
understood the neecis of basic research. T was one of many
visitors who joined in the excitement of the place. In later
years other corporate managers were less supportive, but by
that time Bessel and his group had achieved a position of
strength.
In a former wine cellar Bessel assembled an improved
version of his machine, which now couIc! be called a split-
beam phosphoroscope. A new sector program allowect mea-
surements just before anc! just after the flash and was espe-
cially useful in studying the absorbance changes near 700
nm. Now the characteristics of the "700 pigment" were re-
ve-aled: it was repeatedly bleachect, even by rather weak
flashes of far-rect light, and its absorption regained in each
subsequent dark period. Of course the measuring light (as at
700 nm) was itself a "far-red." If the measuring beam was
macle brighter, there was nothing left for far-red flashes to
bleach. But now a new behavior appeared: if the flashes were
rec! (insteact of far-red), then each flash restored absorption
of the "700 pigment." Here were reciprocal effects of recI and
far-red light like those clescribec! by Robert Emerson for the
quantum yield of oxygen evolution.
At this stage Bessel acquired a young collaborator: George
Hoch, a biochemist. The collaboration doubled the number
of techniques of measurement but far more than cloublect
the imaginative interaction of two minds bouncing ideas back
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134
BIOGRAPHICAL MEMOIRS
anct forth. The first fruit of their effort came in a joint paper
in a 1960 symposium entitled "Light anct I,ife." In that paper
they opened the modern era of thinking about photosyn-
thesis by the first explicit statement that there must be two
photochemical reactions "the first sensitizer! by chlorophyll a
and a direct photochemical bleaching of P700; the second
sensitize<] by accessory pigment, acting indirectly via media-
tion of dark steps, and restoring P700." The hypothesis was
rephrased in other ways, presented as a diagram, and con-
siderect in terms of other relatecl phenomena. Bessel ancI
George had come with the ultimate excitement of science:
they hac! discoverecl and unclerstooct an important truth that
no one else hacl yet seen. But the reception of their paper
was a disappointment. From the record of discussion one
wouicl judge that they had cIropped an egg instead of a bomb.
They hac! made the tactical error of presenting too many
tiara, too many kincis of experiments. Most of the questions
centered on experimental details that now appear trivial.
Unfortunately, publication of the symposium proceedings
was delayed for over a year. By that time important publica-
tions from the laboratories of Lou Duysens anct Robin Hill
had derivecl indepenclently the two-light-reaction hypothesis
from other (lata. And the Hill-Benclall mocle! became the
convenient Z-scheme.
By 1960 the "black box" of photosynthesis had been
opened up, and its bits and pieces were strewn about. The
following clecacle was a time of fitting the pieces together.
Now Bessel's work moved into high gear. It was remarkable
in terms of the number of papers published and the variety
of subjects adciressed. It was even more remarkable in the
number and diversity of collaborators: some came as senior
scientists, some as postdoctoral stuclents, and some as tech-
nicians; some even came from the ranks of high school stu-
dents employee! by the lab each summer. In the midst of this
OCR for page 139
BESSEL KOK
BIBLIOGRAPHY
139
1940
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containing biocolloids. II. Coacervation of gum-arable by
toluidin-blue and the phenomena accompanying the dissolu-
tion of the coacervate. Proc. K. Ned. Akad. Wet., 43:728-31.
1942
With H. P. Wolvekamp, G. P. Baerends, and W. F. H. M. Mom-
maerts. O2 and CO2-binding properties of the blood of the
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Arch. Neerl. Physiol., 26:203 -11.
~ ~ ~ ~ ,
With H. G. Bungenberg de long. Tissues of prismatic cells con-
taining biocolloids. IV. Morphological changes of the complex
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With H. G. Bungenberg de long. Tissues of prismatic celloidin cells
containing biocolloids. VII. Stagnation effects. Proc. K. Ned.
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1948
A critical consideration of the quantum yield of Chlorella photosyn-
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1949
On the interrelation of respiration and photosynthesis in green
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1951
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1952
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1953
With E. C. Wassink and I. L. P. van Oorschot. The efficiency of
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~ J
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1965
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' 1 ~
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