THE PHOTOCHEMICAL REACTION IN PHOTOSYNTHESIS

BY ROBERT EMERSON ANDWILLIAM ARNOLD

(From the Kerckhoff Laboratories of Biology, California Institute of Technology,
Pasadena)

(Accepted for publication, July 13, 1932)

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I

From the experiments of Warburg and Negelein (1923), we know

that the green alga Chlorella pyrenoidosa can reduce one molecule of
carbon dioxide for each four quanta of light absorbed, when conditions
permit maximum efficiency. Chlorophyll is clearly the substance
absorbing the light quanta, so we m a y inquire how much chlorophyll
must be present for the reduction of one molecule of carbon dioxide.
In a preceding paper (1932) we have presented evidence that the
mechanism involved in the photochemical reaction must undergo a
slower reaction, the so called Blackman reaction, before it can again
take part in the photochemical reaction. Let us consider a cell in
flashing light when the dark periods between flashes are so long that
each unit activated in a given flash has time to complete the Black-
man reaction before the next flash. Increasing the intensity of the
flashes should increase the carbon dioxide reduction per flash until
each unit capable of undergoing the photochemical reaction does so
once in each flash. We say then that the photochemical reaction is
saturated with light. The possibility that any unit will undergo the
light reaction more than once in a single flash may be neglected, be-
cause the time required for the completion of the dark reaction is about
0.02 sec. at 25°C., while the duration of a light flash is 10 -5 sec.
We define one unit arbitrarily as the mechanism which must undergo
the photochemical reaction to reduce one molecule of carbon dioxide.
If we can obtain light flashes of sufficient intensity to saturate the pho-
tochemical reaction, then the number of units in a sample of cells will
equal the number of carbon dioxide molecules reduced per flash. The
191

The Journal of General Physiology

192 PHOTOCHEMICAL REACTION IN PtlOTOSYNTItESIS

t o t a l c h l o r o p h y l l c o n t e n t of t h e s a m p l e d i v i d e d b y t h e n u m b e r of
c a r b o n d i o x i d e m o l e c u l e s r e d u c e d p e r flash will g i v e t h e n u m b e r of
c h l o r o p h y l l m o l e c u l e s p e r u n i t , o r p e r m o l e c u l e of c a r b o n d i o x i d e .
T h e m e a s u r e m e n t of t h i s r a t i o w a s t h e o b j e c t i v e of t h e w o r k d e -
s c r i b e d in t h i s p a p e r .

II
Methods of Measurement
Photosynthesis was measured manometrically in the usual way. The cells
M
were suspended in a mixture of 85 parts i~0 potassium bicarbonate and 15 parts

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potassium carbonate. The flashing light was obtained by discharging a 1 or ½
10
microfarad condenser, charged to about 3000 volts, through a neon tube. The
circuit is described by Emerson and Arnold (1932, p. 395). The tube was flashed
twelve or twenty-one times a second.
To obtain flashes of sufficient intensity to saturate the photochemical reaction
we were obliged to concentrate the light with mirrors. The voltages used on the
condenser were already so high that each tube lasted only a short time. A cylin-
drical mirror was made by splitting a glass tube about 2 cm. in diameter and sil-
vering the outside of one half. This mirror was hung just below the neon tube,
and served to concentrate the light on the vessel containing the photosynthesiz-
ing cells. The sides and top of the vessel were also silvered, and all silvered sur-
faces were copper-plated to protect the silver. Using a cell suspension as a photom-
eter, we found that the mirrors increased the light intensity three to four times.
Ordinary incandescent lamps were not adequate to saturate certain samples of
cells with continuous light. We obtained very intense continuous light from 100
watt high temperature projection lamps by adjusting silvered watch-glasses of
appropriate diameter and curvature so that the images of the filaments fell in the
cell suspensions. Even the intensity so obtained was not wholly satisfactory.
The light intensity was varied quantitatively by attaching neutral filters of
known percentage transmission to the bottoms of the vessels. These filters are
sufficiently non-selective for white light, but we found denser filters could transmit
more red light than their indicated values. The filters used in our experiments
with neon light were calibrated for red with the spectrophotometer.
The chlorophyll content of the cells had to be determined in absolute units.
We are much indebted to Dr. Hans Gaffron for a sample of chlorophyll a + b
prepared from Chlorella, with which we standardized our measurements. 13.3
rag. of dry chlorophyll, weighed accurately to 0.1 rag., were dissolved in 1 liter
of pure methanol. The extinction coefficient of this solution was measured with
a K6nig-Martens spectrophotometer, using light of 6598.95 7~ from a neon tube.
Prior to measurement the usual Nicol ocular mounted in the divided circle was
 ROBERT EMIERSON AND WILLIA.M~ARNOLD 193

replaced by the Gauss ocular, which shows the lines of the neon spectrum sepa-
rately when the collimator slit is small. The telescope was adjusted so that the
line 6598.95 A was about centered in the field. Then the collimator slit was
opened until the neighboring lines on either side, 6532.88 and 6678.27 A, were
about to fuse with the center line. Telescope and ocular slit were then adjusted
so that only the line 6598.95 A was visible. This gives strictly monochromatic
light of adequate intensity for such chlorophyll solutions as we prepared.
We have used the definition of the extinction coefficient, ~, given in the Hand-
buch der Physik (1925, p. 189) from the equation
--~d
11=1)<10

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This means that ¢ is the reciprocal of that thickness of medium which will
reduce the light to one-tenth its original intensity.
Our standard chlorophyll solution, 13.3 mg. per liter of methanol, gave a value
of 0.634 for ¢ at 6598.95 2~. Reduced to 10 rag. chlorophyll per liter, ¢ - 0.476.
For determining the chlorophyll content of cells, samples of about 10 c.mm.
were used. After being washed in distilled water, boiling water was poured over
them. They were allowed to stand in this for 2 minutes. The treatment does
not decrease the yield of chlorophyll, and allows quicker completion of the extrac-
tion. The cells were then centrifuged out of distilled water, and extracted with
methanol until they were white. Extracts were made up to 25 c.cm. in volumetric
flasks. The technique of measuring the extinction coefficients was the same as
described above for the standard solution.
If v is the volume in c.mm. of ceils extracted, m the molecular weight of chloro-
phyll, e the standard extinction for 10 mg. of chlorophyll per liter, and ~i the coeffi-
cient for the sample, then the number of mols of chlorophyll per c.mm. of sample
equals:
- -
•1 ) < 25
- -
X- 10
v¢m 1000~
For m we used 906.6, the value given by Willst~tter and Stoll (1913, p. 128)
for the average molecular weight of a mixture of chlorophyll a + b. Even large
changes in the ratio a: b would not alter this value as much as 1 per cent, since
the molecular weights of the two chlorophylls differ by only 14 units, according to
Willst~tter and Stoll.
The chlorophyll concentration per unit volume of cells was varied by growing
cultures over different colors of light. I t has been mentioned (Emerson and Ar-
nold, 1932) that satisfactory changes in the chlorophyll content of Cldordla pyre-
noidosa cannot be effected conveniently by the method of lowered iron concentra-
tion used for C. vulgaris (Emerson, 1929). But cultures of C. pyrenoidosa grown
over mercury luminous tubes contain large amounts of chlorophyll per unit amount
of cells, while similar cultures grown over neon tubes contain about one-fourth as
much chlorophyll. Cells grown over 40 watt incandescent lamps develop an in-
termediate amount of chlorophyll. The chlorophyll concentration produced
194 PIIOTOCHE~IICAL REACTION IN P H O T O S Y N T H E S I S

appears to depend on the intensity of the light and the age of the culture, as well as
on the color of the light. The neon light cultures mature faster than the incan-
descent light cultures, the mercury cultures much more slowly. All cultures were
grown at 20°C.
III
EXPERIMENTAL

The flashing light concentrated by mirrors was barely sufficient to

saturate the photochemical reaction, as shown by Fig. i. Photosyn-

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.t

1/ I ,I , I !
Z0 40 60 80
R~lativo light intensity
Fig. 1. Relative light intensity plotted against photosynthesis per flash, in
arbitrary units. Temperature 25°C.
thesis per flash is plotted against intensity. The shape of the curve
shows t h a t higher intensities would probably increase the yield, though
the maximum seems to have been nearly attained. This conclusion
is in h a r m o n y with a possible theoretical explanation of the process
which we shall propose in the last section. Table I gives the data for
the two experiments incorporated in Fig. 1.
The ratio of chlorophyll content to the m a x i m u m height of Curve 1
gives the ratio of carbon dioxide reduction per flash, to a m o u n t of
 ROBERT EMERSON AND WILLIAM ARNOLD 195

chlorophyll. It is important to know whether this ratio remains

constant for different concentrations of chlorophyll, so we have meas-
ured maximum photosynthesis and chlorophyll content for a number
of cell samples grown so as to have different amounts of chlorophyll.
The results, shown in Columns 4 and 10 of Table II, are plotted in Fig.
2. Column 11 of Table II gives the ratio, p, of chlorophyll: photo-
synthesis per flash at saturation. A straight line is the best fit for

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6--

x •

Q •

g..
ff

}/i 0.5
i
1.0 1.5
i,
C, ¢hlo~hyll con¢c.ntI,ation, mol~ p ~ ¢.mm. x 108
FIo. 2. Chlorophyll concentration in mols per c.mm. of cells plotted against
tools of carbon dioxide reduced per flash of light at saturation. Temperature
25°C.
the points in Fig. 2. The slope of the curve, p, is therefore a constant
for different concentrations of chlorophyll. The value of p obtained
by averaging the last column in Table II, 2480 molecules of chlorophyll
per molecule of carbon dioxide reduced per flash, agrees exactly with
the slope of the line chosen as the best fit for the points in Fig. 2.
Column 8 in Table II gives the values of Q, photosynthesis in con-
tinuous light. It is to be understood that Q does not represent light
saturation in all cases, since we were not able to obtain light of suffi-
196 PHOTOCHEMICAL REACTION IN P H O T O S Y N T H E S I S

cient intensity to saturate certain samples of cells. Nevertheless, it is

interesting to compare the highest obtainable values of Q with those
for maximum photosynthesis per flash. Q is plotted against chlo-
rophyU in Fig. 3. The points are more scattered than in Fig. 2. We
attribute the scatter to varying capacities to carry on the Blackman
reaction among different samples of cells, a factor which would not

TABLE I
Photosynthesis in Flashing Light as a Function of Light Intensity. Data for
Fig. 1

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Capacity of condenser ½ t~fd.
Resistance of charging circuit 3340 ohms.
Twenty-one flashes per sec.
3100 volts at rectifier.
Temperature 24.9°C.

Relative light Ah, per 5 min. per Oxygen per $ mln. Relative rate of
intensity c.mrn, ceils, corrected KO~ per c.mm. cells oxygen production,
for respiration M

First experiment
mm. G.mm.

100 1.16 0.52 0.603 14.50

75 1.18 0.52 0.614 14.75
50 0.99 0.52 0.515 12.40
28 0.68 0.52 0.354 8.50
10 0.25 0.52 0.130 3.15
Second experiment

10 0.27 0.50 0.135 2.79

28 0.84 0.50 0.420 8.60
50 1.10 0.50 0.550 11.20
75 1.35 0.50 0.675 13.85
100 1.35 0.50 0.675 13.85

influence saturation in flashing light because the dark periods were

sufficient for the completion of the Blackman reaction between flashes,
but which would surely affect the balance between photochemical
and Blackman reaction in continuous light. This might also explain
why we could not saturate certain samples with continuous light.
The cells with low chlorophyll content tended to fall short of satura-
tion, even in our most intense continuous light. If cells with high
 ROBERT EMERSON AND WILLIAM ARNOLD 197

chlorophyll content should have their capacity for the Blackman re-
action less well developed in proportion to their chlorophyll content
than the cells with low chlorophyll content, then the cells rich in

TABLE II
Photosynthesis in Flashing and Continuous Light as a Function of Chlorophyll
Content. Data.for Figs. 2 and 3
Capacity of condenser ½ or 1 / g d .
Resistance of charging circuit 3300 to 7500 ohms.
Twelve flashes per sec.
3100 volts a t rectifier.

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T e m p e r a t u r e 25°C.

¢ for i 0
c.mm. P,
cells, Source of l i g h t for
25 cc. culture c
meth- m
anol

0.0816 Neon 2660

0. 204 Mercury 2380
0.146 Neon 2380
0.242 Mercury 2780
0.0672 Neon 2480
0.144 Mercury 2010
0.0544 Neon
0. 250 40 watt lamps
0.0423 Neon
0.224 40 watt lamps
0.118 Neon 2660
0.218 Mercury 3120
0.116 Neon 1980
0.284 Mercury 2310
0.303 Mercury 2460

2480

chlorophyll would be saturated with light at lower intensities than

those poor in chlorophyll. This would explain the direction of curva-
ture of the line drawn through the points in Fig. 3. It would also
explain the shape of the chlorophyll-photosynthesis curves published by
Emerson (1929) for C. vulgaris. Those curves were made with cul-
198 PttOTOCttEMICAL REACTIONIN PHOTOSYNTHESIS

tures of the same age and nearly the same density. The maintain-
ance of equal age and density in cultures of C. pyrenoidosa grown over
mercury and neon lights is not as easily possible, since the cultures
mature so much faster in the neon light. The capacity for the Black-
man reaction may well depend on the age and rate of growth of the
culture, as well as on the chlorophyll content.
We are aware that we have published curves (1932, p. 413, Fig. 12)
indicating identical capacity for the Blackman reaction relative to

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o 4
o
-~ y O • •
0

0
/ .

I
0.4
I
0.8
I
1.~
I
1.6
C, l~Iols chlox~ophyll per c.mm. cells × 108
FIo. 3. Concentration of chlorophyll in tools per c.mm. of cells plotted against
mols of carbon dioxide reduced per second in high intensity continuous light.
Temperature 25°C.

chlorophyll content in two very different samples of ceils, grown over

red and blue light. We feel that we must attribute this result to
chance, because the factors governing the development of the capacity
for the Black.man reaction are evidently not yet under our control.

IV
Theoretical
We can give no adequate interpretation of our ratio of 2480 mole-
cules of chlorophyll per molecule of carbon dioxide reduced per flash.
 ROBERT EMERSON AND WILLIAM ARNOLD 199

Warburg and Negelein (1923) found that under favorable conditions

Chlorella pyrerwidosa could reduce one molecule of carbon dioxide
for every four quanta absorbed. The light emitted by our neon tube
is rich in red, a color strongly absorbed by chlorophyll; and we know
that at high intensities the yield of photosynthesis per unit time of
light is greatly improved by illuminating with short flashes separated
by long dark periods. As yet we know nothing of the quantum effi-
ciency in flashing light. We are forced to conclude that this is very low,
or that most of the chlorophyll is not absorbing light. The fact that
our maximum intensities nearly saturated the photochemical reaction

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does not mean necessarily that every chlorophyll molecule was absorb-
ing light in each flash. We need only suppose that for every 2480
molecules of chlorophyll there is present in the cell one unit capable of
reducing one molecule of carbon dioxide each time it is suitably acti-
vated by light. At our maximum intensity each flash activated nearly
all these units.
It is also possible that the molecular weight of chlorophyll in the
cell may be higher than that of extracted chlorophyll.
Knowing the number of units and the maximum rates of photosyn-
thesis in continuous and flashing light, we can calculate the average
time required for one unit to go through the cycle of photochemical
and Blackman reactions. We suppose the Blackman reaction must
be completed each time the photochemical reaction takes place, be-
fore the unit involved is again free to undergo the photochemical re-
action. Therefore the mean time of one cycle, which we call S, will
be longer at low temperatures than at high ones.
To calculate S, we shall let C be the chlorophyll content of the cells;
m the photosynthesis per flash at saturation; and Q the photosynthesis
per second at saturation with continuous light. C, m, and Q are in
mols per c. mm. of cells. We have used m as a measure of the number
of units capable of undergoing the photochemical reaction. If S is
the mean time required for a unit to undergo photochemical and
Black_man reactions, then m/S is the maximum possible rate of photo-
synthesis in continuous light at saturation, so we may write

For a sample of cells whose values of m and Q fall on Curves 2 and

200 PIIOTOCHEMICAL REACTION IN P H O T O S Y N T H E S I S

3, m, which is independent of temperature, is 2.52 X 10-12 tools. At

25 ° Q is 2.08 X 10-1° mols. From these figures S is equal to 1.2 X
10-2 sec.
We can also estimate the value of S from curves showing the
duration of the dark reaction after a flash of light. The mean time of
one cycle (neglecting the duration of the light reaction because it is
very short compared to that of the dark reaction) will be approxi-
mately equal to the time required for the Blackman reaction, when
taken by itself, to convert half the product of the photochemical reac-
tion. This time can be read from numerous curves for the dark reac-

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tion which we have published previously (1932) for temperatures below
25 °. Fig. 8 (1932, p. 403) shows that at 25 ° the Blackman reac-
tion is substantially completed in less than 0.035 sec., the shortest dark
time used. It would be half completed in about half this time, the
exact point being determined by the order of the Blackman reaction,
which we do not yet know. But we may say from Fig. 8 that S is
about 0.017 sec., a figure of the same order of magnitude as the value
m
calculated from ~? at 25 °. The value of Q for lower temperatures can
be calculated roughly from the known temperature coefficient of pho-
tosynthesis, and the resulting values of S remain of the same order of
magnitude as those estimated from the dark-time curves for corre-
sponding temperatures.
We conclude, then, that at 25 ° the mean time required for a unit
to complete the cycle of photochemical and Blackman reactions, re-
ducing one molecule of carbon dioxide, is somewhere between 0.01
and 0.02 sec.
Now we shall present evidence, derived from the preceding experi-
ments, that the photochemical reaction is of the first order with respect
to light intensity. We will designate the number of units ready to
undergo the photochemical reaction, as they would be after a long dark
period, by N. Under the influence of light some of these units are
activated and become ready to undergo the Blackman reaction. We
will designate the number of these photoactivated units as n. They
are reconverted by the Blackman reaction. We will let K be the
value of N + n. K is a constant for a given sample of cells, and is
proportional to the chlorophyll content of the sample. We assume
 ROBERT EMERSON" AND WILLIAm[ ARNOLD 201

also that the rate, R, at which the units undergo the photochemical
reaction, is proportional to the light intensity and to the value of N.
We need not make any assumptions about the reconversion of the
units as long as we consider photosynthesis in flashing light with long
dark periods.
Our assumptions are:
iv + n = K, (1)

and

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g -~ A I N . (2)

In equation (2) A is the velocity constant for the reaction N +

n. Equation (2) can be expressed in differential form:
dN
-- = -- AIN, (3)
dt

with t representing the time, neglecting the Blackman reaction because

t is short.
Integrating both sides of (3),
-- f Aldt.
N = ce (4)

When t = 0, equation (4) becomes

N,= c,

but we know that after a long dark period, all the units are ready to
undergo the light reaction, so n = 0, and N = K. Hence we m a y re-
place the integration constant c in equation (4) by K. A is a constant,
and may be written before the integral sign:

N = K e- a f 1at. (5)

We have good evidence that E, the total energy in a flash, is equiva-

lent to f I d t. By inserting different sized choke coils in series with
the condenser and the neon tube, the light flashes can be made slower
and less intense, while their energy content, E, remains proportional
to the charge on the condenser, a constant. By means of our spin-
ning thread (Emerson and Arnold, 1932, p. 397) we estimated that our
202 PHOTOCHEMICAL REACTION IN PHOTOSYNTHESIS

largest choke coil increased the duration of the flash about 100 times.
The maximum intensity of the flash is correspondingly decreased since
the total energy liberated is the same. Photosynthesis was measured
in flashes of various duration while E was kept constant. Unfortu-
nately the fact that the energy emitted by the tube remained constant
does not mean that the energy absorbed by the cells was the same for
the various flashes. The wave length distribution of the flashes
differed with the different choke coils. The slower flashes were darker
red, and may have emitted more energy as heat. For this reason
the results are not quantitative. Nevertheless, we were only able to

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find differences of 9 per cent in the photosynthesis per flash, using the
largest choke coil. These experiments indicate that it is correct to use
the first power of the intensity in equation (5). A higher power of I
would require that the amount of photosynthesis per flash change with
changes in intensity and duration of flash produced by the different
choke coils. It appears from the experiments that f I dt, which
remains equal to the charge on the condenser, a constant, produces
the same photosynthesis even when the time and intensity are changed
over a wide range.
We will now let N1 denote the value of N remaining at the end of
a light flash, and rewrite equation (5):
5"1 = K e- a f zet (6)

K - N1 will be the number of units activated, measured by M.

M = K - - N1,

or

M = K - Ke--a'fldt" (7)

Subtracting both sides of equation (7) from K, and simplifying,

K -- M -A f Idt
K
or

log K--M__
- - _A~Idt. (8)
K d
 ROBERT EMERSON AND WILLIAM ARNOLD 203

K--M
According to equation (8), the log of K plotted against f l d t
should give a straight line of slope - A , intersecting the logarithmic
axis where M = 0. This plot can be made from Fig. 1. f l d t is
proportional to the intensity scale, and M is the height of the curve at
any point. K, being proportional to C, the chlorophyll content, is
proportional to the maximum height to which the curve rises. It was
mentioned in the discussion of Fig. 1 that our light intensities did not

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~ -1.5

,
20
,
40 60
, \ ," 80
'B

E, Th~ en¢~g:,y" pet" l:la~h

FIo. 4. See text for explanation. Curve A is made from Fig. 1, taking K as
the maximum height, 14.5, attained by the curve drawn in Fig. 1. Line B is
plotted from Fig. 1 on the assumption that K would finally reach the value 15.5, if
we could obtain flashes of sufficient intensity.

permit the experimental determination of this maximum height. We

may assign to K either the highest value attained by Curve 1, or a
slightly higher value, the probable maximum. Curve A in Fig. 4 is
a plot of I against log (K -- M) when K is given the value 14.5, the
highest level actually attained b y Curve 1. Curve A deviates most
from a straight line in the region where too low a value of K would
make the largest error. If we assign to K the value 15.5 instead of
14.5, we obtain the line B, in Fig. 4, as required by equation (8).
Since 15.5 is probably nearer the maximum height attainable by the
204 PHOTOCHEMICAL REACTION I N PHOTOSYNTHESIS

curve in Fig. 1 than 14.5, we think our interpretation comes close to

fitting the experimental results.
Any interpretation in which light intensity enters this mechanism
at a power higher than the first, fails to explain these results as satis-
factorily. If we retain the concept that there are units in the photo-
synthetic mechanism which go through the photochemical and Black-
man reactions in a cycle, our explanation is a good fit. Future work
may show that we are mistaken in interpreting the behavior of pho-
tosynthesis in flashing light to mean that a cyclical type of reaction is
governing the process. In this case our scheme will have to be

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abandoned. It covers the photochemical reaction only, but gives
promise that it may be extended to include the Blackman reaction as
well. Qualitatively it will explain the shape of the ordinary intensity
curves published by Warburg (1925), van den Honert (1930), van der
Paauw (1932), and others. We suppose that saturation in continuous
light is reached when the photochemical reaction produces its product
as fast as the Blackman reaction can use it. But an exact interpreta-
tion of the balance between the photochemical and Blackman reac-
tions must await a better understanding of the Blackman reaction.

SUMM2ARY

Measurements of photosynthesis were made in continuous and

flashing light of high intensity, using cells varying in chlorophyll con-
tent. The amount of chlorophyll present per molecule of carbon diox-
ide reduced per single flash of light was found to be about 2480 mole-
cules. The length of time required for one unit in the photosynthetic
mechanism to complete the cycle of photochemical and Blackman
reactions was found to be about 0.02 see. at 25°C. The equation
R = A I N was shown to give a good description of the rate of the photo-
chemical reaction, when A is a velocity constant, I the intensity of
light, and N the number of units in the photosynthetic mechanism.
We are greatly indebted to Mr. Erickson and to the Electrical Prod-
ucts Corporation for the large number of tubes which they furnished
us. Our thanks are due especially to Professor R. C. Tolman for
helpful criticism.
 ROBERT EMERSON AND WILLIAM ARNOLD 205

CITATIONS
Emerson, R., Y. Gen. Physiol., 1929, 12, 609.
Emerson, R., and Arnold, W., Y. Gen. Physiol., 1932, 16, 391.
Geiger, H., and Scheel, K., Handbuch der Physik, Berlin, Julius Springer, 1928.
van den Honert, T. H., Rec. tray. bot. n~erl., 1930, 9.7, 149.
van der Paauw, F., The indirect action of external factors on photosynthesis, Doc-
tor's thesis, Amsterdam, 1932.
Warburg, 0., Biochem. Z., 1925, 166, 386.
Warburg, O., and Negelein, E., Z. phys. Chem., 1923, 1069 191.
Willst/itter, R., and Stoll, A., Untersuchungen fiber Chlorophyll, Berlin, Julius
Springer, 1913.

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