Dena Guo

ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

A History of Photosynthesis
Research
Unlike many other famous scientific breakthroughs like the theory of
relativity or the double helix structure of DNA, photosynthesis wasnt
discovered by a single scientist or even a group of scientists working
together. Instead, several scientists, including five Frenchmen, four
Americans, three Germans, and two ancient Greeks, pieced together the
process of photosynthesis over the course of centuries. Though
photosynthesis developed 3.5 billion years ago, exactly how plants acquired
food wasnt well understood until recent times. In fact, the iconic chemical
equation for photosynthesis , 6 CO2 +6 H 2 O+light C 6 H 12 O 6 +6 O 2 +6 H 2 O,
wasnt complete until the mid-1800s! It all started in 350 B.C. when Aristotle
proposed that plants, like animals, require food to survive. 1 Fifty years later,
Theophrastus observed that plants get nourishment through their roots. 1
Subsequent events in photosynthesis research can be traced through the
development of the chemical equation of photosynthesis.

H 2 O food
In the 1600s, Jan Baptist van Helmont, the founder of pneumatic (gas)
chemistry, grew a willow tree in a pot of soil and documented the trees
weight as it grew. At first, the sapling weighed 5 pounds and the soil weighed
200 pounds. After five years, the willow tree's weight shot up to 169 pounds
while the soils weight remained the same. Van Helmont thus concluded that
since he had only watered the willow tree throughout the five years, its

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

increase in weight as it grew more bark, wood, and roots was due to the
water alone.5

plants +injured air dephlogisticated air

Almost two centuries later in the 1770s, Joseph Priestly lived next to a
brewery and became interested in carbon dioxide. In fact, his first paper
described a process to carbonate water.10 Priestly discovered that air "injured"
by a mouse could be later restored by a green plant. 1 This observation meant
that animals must consume something from air that plants replenish. 6 In his
1772 paper On Different Kinds of Air, Priestly described 9 different gasses,
only 3 of which were previously known.13 His paper was significant because
many scientists still believed Aristotles theory of four elements, in which air
was a single, homogenous element. Two years later, Priestly discovered a
new gas after heating red mercuric oxide ( HgO ). He called the gas
"dephlogisticated air.10
Subsequently that year, Antoine Lavoisier named dephlogisticated air
"oxygen" after investigating its properties. One such property unique to
oxygen that Lavoisier described was that when oxygen was blown over a
wooden splint that was glowing red but not burning, the splint immediately
burst back into flame. This phenomenon, now called a "splint test", is still
used today in chemistry to identify oxygen as a product after a chemical
reaction. Drawing on Priestly's research, Lavoisier realized that animals inhale
oxygen and plants exhale oxygen.16

light + plants O2

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

Then in 1779, Jan Ingenhousz placed plants underwater in a

transparent container and observed them in both direct sunlight and
darkness. He noticed that when the plants were placed in sunlight, the
undersides of the plant leaves produced gas bubbles. However, when the
same plants were placed in the dark, they stopped producing gas bubbles.
Ingenhousz collected the unknown gas that the plants produced and
performed a series of tests to identify the gas. The gas passed the splint test,
confirming that green plants produce oxygen bubbles while under sunlight. 7

CO2 +light ( CH 2 O ) +O2

A decade following Ingenhousz's discovery, Andrea Comparetti
observed plant leaf tissues under a microscope and described the green
granules that they contained. He discerned strands of starch ( C6 H 12 O 6 ), a
carbohydrate, within these green granules, but he did not realize the
significance. These green granules were later named chloroplasts. 18
In his 1800 paper The Physiology of Plants, Jean Senebier stated that
light was responsible for the fixation of inorganic carbon dioxide to organic

carbohydrates

( CH 2 O )

, and that oxygen was only released if plants were in

the presence of carbon dioxide.9 Senebier knew that when plants grew, their
carbon content also grew, and he discovered that the plants increase in
carbon content came from the intake of carbon dioxide. Furthermore, for
every carbon dioxide molecule that the plant absorbed, it released a
molecule of oxygen. He therefore concluded that during photosynthesis,
carbon dioxide is decomposed and the carbon is stored in the plant as a

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

carbohydrate, while oxygen is released. (We now know that he was wrong
because the molecules that assemble the carbohydrate do not come from the
decomposition of carbon dioxide.)12

CO2 + H 2 O+ light ( C H 2 O ) +O2

Four years later, Nicolas-Theodore de Saussure realized that the carbon
dioxide in the atmosphere couldn't fully account for the increase in the plant's
weight as it matured. So, some of the plant's weight had to come from the
water it absorbed.1 In his 1804 paper Chemical Research on Vegetation, he
also proved that plants absorb water and carbon dioxide in sunlight, and that
the increase in a plants weight was also dependent on the absorption of
nitrogen from the soil.15
In 1818, Pierre Joseph Pelletier and Joseph Bienaime Caventou isolated
and named chlorophyll, a pigment in plants that give them a green color.
(Interestingly, they also discovered caffeine.) 17
Following Pelletier and Caventou's research, Henri Dutrochet realized in
1837 that chlorophyll was essential for a plant's ability to use carbon dioxide
to produce starch. He also observed and named the process of osmosis and
deduced that osmosis is used for food transport in plants. 4
Soon after, Matthias Schleiden studied plant structure under a
microscope and hypothesized that every plant was composed of a basic unit,
which he called the cell.14 He additionally proposed that a water molecule is
split into hydrogen and oxygen during photosynthesis, a process now called
hydrolysis.1

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

Then in 1845, Julius Robert von Mayer conjectured that photosynthesis

converted the sun's light energy into chemical energy. Moreover, he believed
that all living organisms depended on the sun's energy to survive. 1

6 CO2 +6 H 2 O+ light C 6 H 12 O6+ 6 O2+6 H 2 O

In 1864, Julius von Sachs demonstrated that starch formed and was
confined in chloroplasts, which agreed with Comparetti's earlier observation
of starch within chloroplasts. Thus, the unknown carbohydrate was solved. He
furthermore demonstrated that the creation of starch in chloroplasts was
dependent on light.11
That same year, Jean Baptiste Boussingault derived the modern
chemical equation for photosynthesis after making a series of specific
measurements.8 When Boussingault published the chemical equation for
photosynthesis, the process was called assimilation. Unfortunately,
assimilation was also used to describe metabolism in animals. In 1893,
Charles Barnes suggested that assimilation in plants should be either called
photosynthesis or photosyntax. Though he personally liked the word
photosyntax, most scientists preferred using photosynthesis, so
photosynthesis was adopted.3
Half a century later in 1941, Cornelis van Niel published the results of
his studies on Chromatium (purple sulfur bacteria) and Thiosarcina rosea
(green sulfur bacteria). He observed that both bacteria photosynthesized, but
they required hydrogen sulfide ( H 2 S ) for photosynthesis instead of oxygen

( O 2 . Van Niel suggested by analogy that the oxygen produced by plants

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

during photosynthesis comes from water ( H 2 O ) instead of carbon dioxide (

CO2 ).2
Later that year, Samuel Ruben and Martin Kamen confirmed van Niels
hypothesis that the oxygen produced in photosynthesis was derived from
water. In their experiment they watered plants with heavy water, or water
containing the

18

O isotope instead of the lighter and more common

16

isotope, and observed that the oxygen emitted by the plants was also

18

O.

When they repeated the experiment with heavy carbon dioxide, the oxygen
emitted by the plants was the normal

16

O. Thus, the emitted oxygen was

clearly derived from water and not carbon dioxide. 12

In 1957, Robert Emerson experimented with the wavelength of light
received by the algae Chlorella and measured the amount of oxygen
produced. He found that while a significant amount of oxygen was produced
with blue light alone, red light alone produced little oxygen. In addition, when
both blue and red light was shone, the amount of oxygen produced far
exceeded the previous amounts. Emerson concluded that there were two
light reactions in photosynthesis: one driven by short-wavelength light (blue
light), called PS2, and one driven by long-wavelength light (red light) called
PS1.16
Though it's fun to imagine a scientist having a Eureka! moment and
discovering something significant, scientific breakthroughs often take the
collaboration of several scientists over the course of years. In fact, it took
over 300 years for scientists to postulate and confirm the chemical equation
for photosynthesis that all biology students take for granted now. And that

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

doesnt even cover subsequent discoveries that add to the understanding of

photosynthesis, like the Calvin cycle (discovered by Melvin Calvin and Andrew
Benson) and the electron transport chain (discovered by Rudolph Marcus). 1
Photosynthesis research is still going on today! This just goes to show that all
science, even cold, hard, facts that are taught in science classes, are
constantly evolving.

Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

Bibliography

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Nov. 2012.
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2012.
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Dena Guo
ENGL 1111 Lerner

November 8, 2012
History of Scientific Concept

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