PHOTOSYNTHESIS TIMELINES

Thomas Brennan
Department of Biology
Dickinson College, Carlisle, PA 17013
brennant@dickinson.edu
c350 BC Aristotle proposes that plants, like animals, require food. Anticipates
Priestley's work 2000 years later by asserting that plants do not require
animals but animals require plants.

c300 BC Theophrastus writes that plants obtain their nourishment through

the roots.
1450 Nicholas of Cusa proposes (but apparently never performs) an
experiment in which a plant is weighed and then planted in a container
containing a weighed amount of soil. After a period of growth, the final
weights of plant and soil, as well as the total weight of water applied, are
determined and compared to the initial values. He speculates this will
demonstrate that the mass of the plant was derived from water rather than
soil.
1648 Jean Baptiste van Helmont performs the experiment proposed by
Nicholas of Cusa nearly 200 years earlier. He concludes that the entire mass
of the plant came from water, but ignores a very slight decrease in the
weight of the soil.
1679 Edme Mariotte proposes that plants obtain part of their nourishment
from the atmosphere.
1727 Stephen Hales writes that plant leaves "very probably" take in
nourishment from the air, and that light may also be involved.
1754 Charles Bonnet observes the emission of gas bubbles by a submerged
illuminated leaf.
1771 Joseph Priestley finds that air which has been made "noxious" by the
breathing of animals or burning of a candle can be restored (i.e., made to
support breathing or combustion again) by the presence of a green plant. He
isolates the gas later identified as oxygen.
1774 Antoine Lavoisier begins to investigate and later names oxygen. He
recognizes that it is consumed in both animal respiration and combustion.
His work discredits the theory of "phlogiston," a hypothetical substance then
believed to be emitted during respiration or combustion, and lays the
foundations of modern chemistry.
1779 Jan Ingenhousz discovers that only the green parts of plants release
oxygen and that this occurs only when they are illuminated by sunlight.

1782 Jean Senebier demonstrates that green plants take in carbon dioxide
from the air and emit oxygen under the influence of sunlight.
1791 Comparetti observes green granules in plant tissues, later identified as
chloroplasts.
1804 Nicolas de Saussure shows that the carbon assimilated from
atmospheric carbon dioxide cannot fully account for the increase in dry
weight of a plant. He hypothesized that the additional weight was derived
from water. At this point, therefore, the basic equation of photosynthesis was
established. It was understood as a process in which a green plant
illuminated by sunlight takes in carbon dioxide and water and converts them
into organic material and oxygen.
1818 Pierre Joseph Pelletier and Joseph Bienaime Caventou give the name
"chlorophyll" to the green pigment in plants.
1837 Rene Dutrochet makes the connection between chlorophyll and the
ability of plants to assimilate carbon dioxide. Also identifies stomata on leaf
surfaces.
1842 Matthias Schleiden postulates that the water molecule is split during
photosynthesis.
1844 Hugo von Mohl makes detailed observations of the structure of
chloroplasts.
1845 Julius Robert von Mayer proposes that the sun is the ultimate source of
energy utilized by living organisms, and introduces the concept that
photosynthesis is a conversion of light energy into chemical energy.
1862 Julius von Sachs demonstrates light-dependent starch formation in
chloroplasts.

1864 Jean Baptiste Boussingault makes accurate quantitative measurements

of carbon dioxide uptake and oxygen production, a step leading to a
balanced equation for photosynthesis: 6CO2 + 12H2O + light energy ---->
C6H12O6 + 6O2 + 6H2O
1873 Emil Godlewski confirms that atmospheric carbon dioxide is the source
of carbon in photosynthesis by showing that starch formation in illuminated
leaves depends upon the presence of carbon dioxide.
1883 Theodor Wilhelm Engelmann illuminates a filamentous alga with light
dispersed through a prism. He finds that motile aerobic bacteria congregate
near the portions illuminated by red and blue wavelengths, thus producing
the first action spectrum for photosynthetic oxygen evolution.
1883 Arthur Meyer describes the chloroplast grana.

1893 Charles Barnes suggests that the process by which illuminated green
plants manufacture carbon compounds be called either "photosyntax" or
"photosynthesis." Although Barnes prefers the former, "photosynthesis" is
adopted into common usage.
1905 F. F. Blackman develops the concept of limiting factors, showing that
photosynthesis consists of two types of reactions: a rapid light-dependent
photochemical process and a slower temperature-dependent biochemical
process. These are later termed "light reactions" and "dark reactions,"
respectively.
1913 Richard Willstatter and Arthur Stoll publish studies on the structure and
chemistry of chlorophyll. Willstatter awarded Nobel Prize, 1915.
1937 Robert (Robin) Hill demonstrates that in the presence of an artificial
electron acceptor isolated chloroplasts can evolve oxygen in the absence of
carbon dioxide.
1941 Cornelis van Niel publishes a summary of his work showing that
photosynthetic bacteria which use H2S as an electron donor produce
elemental sulfur instead of oxygen. He suggests by analogy that the
O2released in plant photosynthesis is derived from H2O rather than CO2.
1941 Samuel Ruben and Martin Kaman use water labeled with the heavy
isotope 18O to confirm that the oxygen produced in photosynthesis comes
from H2O.
1954 Daniel Arnon demonstrates light-dependent ATP formation in
chloroplasts.
1955 Daniel Arnon demonstrates that isolated chloroplasts are capable of
carrying out complete photosynthesis.
1956 Melvin Calvin and coworkers use radioactively labeled 14CO2 to
elucidate the pathway of carbon assimilation in photosynthesis. Calvin
awarded Nobel Prize in 1961.
1957 Robert Emerson describes the "red drop" and "enhancement" effects,
the first indication that the light reactions of photosynthesis consist of two
separate photochemical systems.
1960 Robert Woodward synthesizes chlorophyll. Awarded Nobel Prize, 1965.
1960 Robin Hill and Fay Bendall, based on the work of Emerson and others,
propose the "Z scheme" model for the photosynthetic light reactions.
According to this model, the light reactions consist of two separate
photosystems operating in tandem, each activated by slightly different
wavelengths of light. One photosystem oxidizes water and reduces

cytochrome f, while the other oxidizes cytochrome f and reduces NADP+.

1961 Louis Duysens provides evidence in support of the Z scheme by
demonstrating that exposure to alternating wavelengths of light causes
cytochrome f to switch between oxidized and reduced states.
1968 Roderick Clayton isolates reaction center complexes.
1970 Bessel Kok proposes the "S-states" model of charge accumulation to
explain the stepwise oxidation of H2O and release of O2.
1984 Hans Deisenhofer, Hartmut Michel, and Robert Huber crystallize the
photosynthetic reaction center from a purple bacterium and use X-ray
diffraction techniques to determine its detailed structure. The three share
Nobel Prize, 1988.
2006 Yano, Vittal Yachandra, and co-workers determine the structure of the
manganese-calcium water-splitting complex of Photosystem II.
5/15/08
Photosynthesis is a very important and complex process in nature and some
of its phases are still not completely understood.
Photosynthesis in plants and a few bacteria is responsible for feeding nearly
all life on Earth. It does this by taking energy from the sun and converting it
into a storable form, usually glucose, which plants use for their own life
processes. Animals that consume plants also make use of this energy, as do
those that consume those that consume plants, and so on to the top of the
food chain.
As important a job as making all of the world's food is, there's another vital
function that photosynthesis performs: It generates the oxygen that oxygenbreathing animals need to survive. But here we animals repay the favor. We
exhale the carbon dioxide that plants need for photosynthesis.
People have long been interested in how plants obtain the nutrients they
use for growth. The early Greek philosophers believed that plants obtained
all of their nutrients from the soil. This was a common belief for many
centuries.
In the first half of the seventeenth century, Jan Baptista van Helmont (15791644), a Dutch physician, chemist, and alchemist, performed important
experiments which disproved this early view of photosynthesis. He grew a
willow tree weighing 5 lb (2.5 kg) in a clay pot which had 200 lb (91 kg) of
soil. Five years later, after watering his willow tree as needed, it weighed
about 169 lb (76.5 kg) even though the soil in the pot lost only 2 oz (56 g) in
weight. Van Helmont concluded that the tree gained weight from the water
he added to the soil, and not from the soil itself. Although van Helmont did
not understand the role of sunlight and atmospheric gases in plant growth,
his early experiment advanced our understanding of photosynthesis.
In 1771, the noted English chemist Joseph Priestley performed a series of
important experiments which implicated atmospheric gases in plant growth.
Priestley and his contemporaries believed a noxious substance, which they

called phlogiston, was released into the air when a flame burned. When
Priestley burned a candle within an enclosed container until the flame went
out, he found that a mouse could not survive in the "phlogistated" air of the
container. However, when he placed a sprig of mint in the container after the
flame had gone out, he found that a mouse could survive. Priestley
concluded that the sprig of mint chemically altered the air by removing the
"phlogiston." Shortly after Priestly's experiments, Dutch physician Jan
Ingenhousz (1730-1799) demonstrated that plants "dephlogistate" the air
only in sunlight, and not in darkness. Further, Ingenhousz demonstrated that
the green parts of plants are necessary for" dephlogistation" and that
sunlight by itself is ineffective. As Ingenhousz was performing his
experiments, the celebrated French chemist Antoine Lavoisier (1743-1794)
disproved the phlogiston theory. He conclusively demonstrated that candles
and animals both consume a gas in the air which he named oxygen. This
implied that the plants in Priestley's and Ingenhousz's experiments produced
oxygen when illuminated by sunlight. Considered by many as the founder of
modern chemistry, Lavoisier was condemned to death and beheaded during
the French revolution. Lavoisier's experiments stimulated Ingenhousz to
reinterpret his earlier studies of "dephlogistation." Following Lavoisier,
Ingenhousz hypothesized that plants use sunlight to split carbon dioxide
(CO2) and use its carbon (C) for growth while expelling its oxygen (O2) as
waste. This model of photosynthesis was an improvement over Priestley's,
but was not entirely accurate.
Ingenhousz's hypothesis that photosynthesis produces oxygen by splitting
carbon dioxide was refuted about 150 years later by the Dutch-born
microbiologist Cornelius van Niel (1897-1985) in America. Van Niel studied
photosynthesis in anaerobic bacteria, rather than in higher plants. Like
higher plants, these bacteria make carbohydrates during photosynthesis.
Unlike plants, they do not produce oxygen during photosynthesis and they
use bacteriochlorophyll rather than chlorophyll as a photosynthetic pigment.
Van Niel found that all species of photosynthetic bacteria which he studied
required an oxidizable substrate. For example, the purple sulfur bacteria use
hydrogen sulfide as an oxidizable substrate and the overall equation for
photosynthesis in these bacteria is: On the basis of his studies with
photosynthetic bacteria, van Niel proposed that the oxygen which plants
produce during photosynthesis is derived from water, not from carbon
dioxide. In the following years, this hypothesis has proven true. Van Niel's
brilliant insight was a major contribution to our modern understanding of
photosynthesis. The study of photosynthesis is currently a very active area of
research in biology. Hartmut Michel and Johann Deisenhofer recently made
a very important contribution to our understanding of photosynthesis. They
made crystals of the photosynthetic reaction center fromRhodopseudomonas
viridis, an anaerobic photosynthetic bacterium, and then used x-ray
crystallography to determine its three-dimensional structure. In 1988, they
shared the Nobel Prize in Chemistry with Robert Huber for this groundbreaking research. Modern plant physiologists commonly think of
photosynthesis as consisting of two separate series of interconnected
biochemical reactions, the light reactions and the dark reactions. The light
reactions use the light energy absorbed by chlorophyll to synthesize labile
high energy molecules. The dark reactions use these labile high energy
molecules to synthesize carbohydrates, a stable form of chemical energy
which can be stored by plants. Although the dark reactions do not require
light, they often occur in the light because they are dependent upon the light
reactions. In higher plants and algae, the light and dark reactions of

photosynthesis occur in chloroplasts, specialized chlorophyll-containing

intracellular structures which are enclosed by double membranes.