Evolution

Main article: Evolution of photosynthesis

Life timeline

This box: 
 view
 talk
 edit
Earth formed
−4500 —   ← (4540 mya)
← Earliest water
– Water ← Earliest life
—   ← LHB meteorites
← Earliest oxygen
– Single-celled life ← Pongola glaciation*
−4000 — ← Atmospheric oxygen
  ← Huronian glaciation*
– ← Earliest multicellular life
Photosynthesis
← Earliest fungi
—
  ← Sexual reproduction
– ← Earliest plants
Eukaryotes ← Earliest animals
−3500 — ← Cryogenian ice age*
  ← Ediacaran biota
–
← Cambrian explosion
Multicellular life
— ← Andean glaciation*
  ← Earliest tetrapods
– ← Karoo ice age*
P ← Earliest apes / humans
−3000 — l ← Quaternary ice age*
– a
n
— t
s
–
 
−2500 —
Arthropods Molluscs
–
Flowers
—
Dinosaurs
–
 
−2000 —
Mammals
–
Birds
—
Primates
–
H
−1500 — a
d
– e
— a
n
–
 −1000 —
A
– r
c
— h
e
–
a
−500 — n

0 —
P
r
o
t
e
r
o
z
o
i
c

P
h
a
n
e
r
o
z
o
i
c

 
 
(million years ago)

*Ice Ages

Early photosynthetic systems, such as those in green and purple sulfur and green and purple

nonsulfur bacteria, are thought to have been anoxygenic, and used various other molecules than
water as electron donors. Green and purple sulfur bacteria are thought to have
used hydrogen and sulfur as electron donors. Green nonsulfur bacteria used various amino and
other organic acids as an electron donor. Purple nonsulfur bacteria used a variety of nonspecific
organic molecules. The use of these molecules is consistent with the geological evidence that
Earth's early atmosphere was highly reducing at that time.[56]
Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4
billion years old.[57][58] More recent studies, reported in March 2018, also suggest that
photosynthesis may have begun about 3.4 billion years ago. [59][60]
The main source of oxygen in the Earth's atmosphere derives from oxygenic photosynthesis, and
its first appearance is sometimes referred to as the oxygen catastrophe. Geological evidence
suggests that oxygenic photosynthesis, such as that in cyanobacteria, became important during
the Paleoproterozoic era around 2 billion years ago. Modern photosynthesis in plants and most
photosynthetic prokaryotes is oxygenic. Oxygenic photosynthesis uses water as an electron
donor, which is oxidized to molecular oxygen (O
2) in the photosynthetic reaction center.

Symbiosis and the origin of chloroplasts

Plant cells with visible chloroplasts (from a moss, Plagiomnium affine)

Several groups of animals have formed symbiotic relationships with photosynthetic algae. These

are most common in corals, sponges and sea anemones. It is presumed that this is due to the
particularly simple body plans and large surface areas of these animals compared to their
volumes.[61] In addition, a few marine mollusks Elysia viridis and Elysia chlorotica also maintain a
symbiotic relationship with chloroplasts they capture from the algae in their diet and then store in
their bodies (see Kleptoplasty). This allows the mollusks to survive solely by photosynthesis for
several months at a time.[62][63] Some of the genes from the plant cell nucleus have even been
transferred to the slugs, so that the chloroplasts can be supplied with proteins that they need to
survive.[64]
An even closer form of symbiosis may explain the origin of chloroplasts. Chloroplasts have many
similarities with photosynthetic bacteria, including a circular chromosome, prokaryotic-
type ribosome, and similar proteins in the photosynthetic reaction center. [65][66] The endosymbiotic
theory suggests that photosynthetic bacteria were acquired (by endocytosis) by
early eukaryotic cells to form the first plant cells. Therefore, chloroplasts may be photosynthetic
bacteria that adapted to life inside plant cells. Like mitochondria, chloroplasts possess their own
DNA, separate from the nuclear DNA of their plant host cells and the genes in this chloroplast
DNA resemble those found in cyanobacteria.[67] DNA in chloroplasts codes for redox proteins
such as those found in the photosynthetic reaction centers. The CoRR Hypothesis proposes that
this co-location of genes with their gene products is required for redox regulation of gene
expression, and accounts for the persistence of DNA in bioenergetic organelles. [68]

Photosynthetic eukaryotic lineages

Symbiotic and kleptoplastic organisms excluded:

 The glaucophytes and the red and green algae—clade Archaeplastida (unicellular and

multicellular)
 The cryptophytes—clade Cryptista (unicellular)
 The haptophytes—clade Haptista (unicellular)
 The dinoflagellates and chromerids in the superphylum Myzozoa—
clade Alveolata (unicellular)
 The ochrophytes—clade Heterokonta (unicellular and multicellular)
 The chlorarachniophytes and three species of Paulinella in the phylum Cercozoa—
clade Rhizaria (unicellular)
 The euglenids—clade Excavata (unicellular)
Except for the euglenids, which is found within the Excavata, all of them belong to
the Diaphoretickes. Archaeplastida and the photosynthetic Paulinella got their plastids— which
are surrounded by two membranes, through primary endosymbiosis in two separate events by
engulfing a cyanobacterium. The plastids in all the other groups have either a red or green algal
origin, and are referred to as the "red lineages" and the "green lineages". In dinoflaggelates and
euglenids the plastids are surrounded by three membranes, and in the remaining lines by four.
A nucleomorph, remnants of the original algal nucleus located between the inner and outer
membranes of the plastid, is present in the cryptophytes (from a red algae) and
chlorarachniophytes (from a green algae).[69] Some dinoflaggelates which have lost their
photosyntethic ability have later regained it again through new endosymbiotic events with
different algae. While able to perform photosynthesis, many of these eukaryotic groups
are mixotrophs and practice heterotrophy to various degrees.

Cyanobacteria and the evolution of photosynthesis

The biochemical capacity to use water as the source for electrons in photosynthesis evolved
once, in a common ancestor of extant cyanobacteria (formerly called blue-green algae), which
are the only prokaryotes performing oxygenic photosynthesis. The geological record indicates
that this transforming event took place early in Earth's history, at least 2450–2320 million years
ago (Ma), and, it is speculated, much earlier. [70][71] Because the Earth's atmosphere contained
almost no oxygen during the estimated development of photosynthesis, it is believed that the first
photosynthetic cyanobacteria did not generate oxygen. [72] Available evidence from geobiological
studies of Archean (>2500 Ma) sedimentary rocks indicates that life existed 3500 Ma, but the
question of when oxygenic photosynthesis evolved is still unanswered. A clear paleontological
window on cyanobacterial evolution opened about 2000 Ma, revealing an already-diverse biota of
Cyanobacteria. Cyanobacteria remained the principal primary producers of oxygen throughout
the Proterozoic Eon (2500–543 Ma), in part because the redox structure of the oceans favored
photoautotrophs capable of nitrogen fixation.[citation needed] Green algae joined cyanobacteria as the
major primary producers of oxygen on continental shelves near the end of the Proterozoic, but it
was only with the Mesozoic (251–66 Ma) radiations of dinoflagellates, coccolithophorids, and
diatoms did the primary production of oxygen in marine shelf waters take modern form.
Cyanobacteria remain critical to marine ecosystems as primary producers of oxygen in oceanic
gyres, as agents of biological nitrogen fixation, and, in modified form, as the plastids of marine
algae.[73]

Experimental history
Discovery
Although some of the steps in photosynthesis are still not completely understood, the overall
photosynthetic equation has been known since the 19th century.

Portrait of Jan Baptist van Helmont by Mary Beale, c.1674

Jan van Helmont began the research of the process in the mid-17th century when he carefully
measured the mass of the soil used by a plant and the mass of the plant as it grew. After noticing
that the soil mass changed very little, he hypothesized that the mass of the growing plant must
come from the water, the only substance he added to the potted plant. His hypothesis was
partially accurate – much of the gained mass also comes from carbon dioxide as well as water.
However, this was a signaling point to the idea that the bulk of a plant's biomass comes from the
inputs of photosynthesis, not the soil itself.
Joseph Priestley, a chemist and minister, discovered that when he isolated a volume of air under
an inverted jar and burned a candle in it (which gave off CO 2), the candle would burn out very
quickly, much before it ran out of wax. He further discovered that a mouse could
similarly "injure" air. He then showed that the air that had been "injured" by the candle and the
mouse could be restored by a plant.[74]
In 1779, Jan Ingenhousz repeated Priestley's experiments. He discovered that it was the
influence of sunlight on the plant that could cause it to revive a mouse in a matter of hours. [74][75]
In 1796, Jean Senebier, a Swiss pastor, botanist, and naturalist, demonstrated that green plants
consume carbon dioxide and release oxygen under the influence of light. Soon
afterward, Nicolas-Théodore de Saussure showed that the increase in mass of the plant as it
grows could not be due only to uptake of CO2 but also to the incorporation of water. Thus, the
basic reaction by which photosynthesis is used to produce food (such as glucose) was outlined. [76]

Refinements
Cornelis Van Niel made key discoveries explaining the chemistry of photosynthesis. By studying
purple sulfur bacteria and green bacteria he was the first to demonstrate that photosynthesis is a
light-dependent redox reaction, in which hydrogen reduces (donates its – electron to) carbon
dioxide.
Robert Emerson discovered two light reactions by testing plant productivity using different
wavelengths of light. With the red alone, the light reactions were suppressed. When blue and red
were combined, the output was much more substantial. Thus, there were two photosystems, one
absorbing up to 600 nm wavelengths, the other up to 700 nm. The former is known as PSII, the
latter is PSI. PSI contains only chlorophyll "a", PSII contains primarily chlorophyll "a" with most of
the available chlorophyll "b", among other pigments. These include phycobilins, which are the red
and blue pigments of red and blue algae respectively, and fucoxanthol for brown algae and
diatoms. The process is most productive when the absorption of quanta are equal in both the
PSII and PSI, assuring that input energy from the antenna complex is divided between the PSI
and PSII system, which in turn powers the photochemistry. [13]
Robert Hill thought that a complex of reactions consisted of an intermediate to cytochrome
b6 (now a plastoquinone), and that another was from cytochrome f to a step in the carbohydrate-
generating mechanisms. These are linked by plastoquinone, which does require energy to
reduce cytochrome f for it is a sufficient reductant. Further experiments to prove that the oxygen
developed during the photosynthesis of green plants came from water, were performed by Hill in
1937 and 1939. He showed that isolated chloroplasts give off oxygen in the presence of
unnatural reducing agents like iron oxalate, ferricyanide or benzoquinone after exposure to light.
The Hill reaction[77] is as follows:
2 H2O + 2 A + (light, chloroplasts) → 2 AH2 + O2
where A is the electron acceptor. Therefore, in light, the electron acceptor is reduced and
oxygen is evolved.
Samuel Ruben and Martin Kamen used radioactive isotopes to determine that the oxygen
liberated in photosynthesis came from the water.
Melvin Calvin works in his photosynthesis laboratory.

Melvin Calvin and Andrew Benson, along with James Bassham, elucidated the path of

carbon assimilation (the photosynthetic carbon reduction cycle) in plants. The carbon
reduction cycle is known as the Calvin cycle, which ignores the contribution of Bassham and
Benson. Many scientists refer to the cycle as the Calvin-Benson Cycle, Benson-Calvin, and
some even call it the Calvin-Benson-Bassham (or CBB) Cycle.
Nobel Prize-winning scientist Rudolph A. Marcus was later able to discover the function and
significance of the electron transport chain.
Otto Heinrich Warburg and Dean Burk discovered the I-quantum photosynthesis reaction
that splits the CO2, activated by the respiration. [78]
In 1950, first experimental evidence for the existence of photophosphorylation in vivo was
presented by Otto Kandler using intact Chlorella cells and interpreting his findings as light-
dependent ATP formation.[79] In 1954, Daniel I. Arnon et al. discovered
photophosphorylation in vitro in isolated chloroplasts with the help of P32.[80][81]
Louis N.M. Duysens and Jan Amesz discovered that chlorophyll "a" will absorb one light,
oxidize cytochrome f, while chlorophyll "a" (and other pigments) will absorb another light but
will reduce this same oxidized cytochrome, stating the two light reactions are in series.

Development of the concept

In 1893, Charles Reid Barnes proposed two terms, photosyntax and photosynthesis, for the
biological process of synthesis of complex carbon compounds out of carbonic acid, in the
presence of chlorophyll, under the influence of light. Over time, the
term photosynthesis came into common usage as the term of choice. Later discovery of
anoxygenic photosynthetic bacteria and photophosphorylation necessitated redefinition of
the term