UNIT IV: PHOTOSYNTHESIS

OVERVIEW: RESEARCH MILESTONES IN PLANT

PHYSIOLOGY
The Process That Feeds the Biosphere
Sir Francis Bacon
Photosynthesis
✓ Published one of the first plant physiology
The process converts solar energy into chemical energy. ✓ Experiments in 1627 in the book, Sylva
Sylvarum
➢ Directly or in directly, photosynthesis
nourishes almost the entire living world Jan Baptist van Helmont
AUTOTROPHS ➢ Published what is considered the first
quantitative experiment in plant
organism that are capable of synthesizing its own food
physiology in 1648.
➢ Producers of the biosphere, producing
John Woodward
organic molecules from CO2 and other
inorganic molecules. ➢ In 1699, he published experiments on
➢ Example: growth of spearmint in different sources of
Plants, multicellular alga, cyanobacteria, water.
unicellular protest, and purple sulfur
bacteria Stephen Hales

➢ Considered the Father of Plant

Physiology for the many experiments in
the 1727.

Julius van Sachs

➢ Unified the pieces of plant physiology and

put them together as a discipline. His
Lehbuch der Botanik was the plant
physiology bible of its time.

Researchers discovered in the 1800s that plants absorb

essential mineral nutrients as inorganic ions in water.

HISTORY OF PHOTOSYNTHESIS

320 B-C – Aristotle and Theoprastus

➢ Thought that plants drive all their

nourishment directly from soil.

1721- S. HALES – light and air are the source of

HETEROTROPHS nutrition for leaves
organisms that cannot manufacture its own food 1772- JOSEPH PRISTLEY – evolution of oxygen by
plants, showed oxygen is consumed by animals.
✓ Consumers- of the biosphere, including
humans, depend on photoautotrophs for 1779- J. INGEN-HOUSZ- solar radiation and green
food and O2 leaf pigment are indispensable for gas exchange (O2 and
CO2)

1782- JEAN SENEBIAR – rate of purification of air is

carried out by green plants only and that light is essential
to the process.

1804- T de Saussure- water is indispensable for CO2,

assimilation the ratio between released O2 and absorbed
CO2 is equal to 1.

1837- Durachet – established connection between

photosynthesis and chlorophyll.

1842- J.R. Mayer- formulated the principal law of

energy transformation in photosynthesis.

1845- Leibig – organic matter synthesized during

photosynthesis is derived from carbon dioxide and water.
1882- T.W. Engelmann – obtained the first
photosynthetic action spectrum with red and blue zone
maxima.

1882- Sachs – proved that strach is synthesized by plans

in light dependent reaction.

1905- F.F. Blackman & G.L.C. Matthaei – use of light

in photosynthesis is dependent an CO2 supply

1913- Wilstaller and Stoll – isolated chlorophyll and

later determine its structure.

1919- O. Warburg- found that the efficiency of

photosythesis is higher with intermittent light

1922- O. Warburg and E. Negelein – determie precise PHOTOSYNTHESIS

spectra of photosynthesis efficiency.
The process by which autotrophic organisms use
1923- Thunberg – recognized that CO2 is reduced and light energy to make sugar and oxygen gas from
water oxidized during photosynthesis. carbon dioxide and water
1932- R. Emerson and W. Arnold – existence of the
photosynthesis (chlorophyll) unit based on experiments
with intermittent illumination.

1936- Wood and Werkman- Distinguished between the

light and dark reaction.

1939- R. Hill – measured O2 production in isolated

chloroplast after the addition of an electron acceptor.

1940- S. Ruben and A.P. Vinogrdov – oxygen released

in photosynthesis originated from water.

1948- Calvin and Benson – phospoglycerate is an early

product of carbon dioxide fixation.

1951- A.H. Mehler- oxygen may serve as an acceptor of 2 stages:

electrins
1) Light Dependent Process (Light reaction)
1954- D.I. Arnon, M.B. Allen and F.R. Whatley- 2) Light Independent Process/The Calvin
chloroplast is the site of CO2 fixation and
cycle (Dark Reaction)
phosporylation. 1) Light reactions convert solar energy to
1956- Calvin, Bassham and Benson – explanation of chemical energy
photosynthetic fixation of CO2. o Occur grana
o Water is split, there is release of oxygen.
CHLOROPLASTS: THE SITE OF o Generate ATP from ADP by
PHOTOSYNTHESIS IN PLANTS photophosphorylation
o Reduce NADP+ to NADPH is formed.
➢ The green color is from cholorophyll, the 2) Calvin cycle makes sugar from carbon dioxide
green pigment with chloroplasts o Occurs in the stroma
➢ Light energy absorbed by cholorphyll o ATP provides the energy for sugar
drives the synthesis of organc molecules in synthesis
the chloroplast o NADPH provides the electrons for the
➢ CO2 enters and O2 exits the leaf through (may na unang word kaso di malabo
microscopic pores called stomata sakin) of carbon dioxide to glucose.
➢ Found in the mesophyll cells, interior
tissue of the leaf (30-40 chloroplasts). THE NATURE OF SUNLIGHT
➢ Chlorophyll is in the membranes of
thylakoids (connected sacs in the ✓ Light is a form of electromagnetic energy also
chloroplast) called electromagnetic radiation
- Thylakoids may be ✓ Wavelength is the distance betweem crests of
stacked in columns called waves
grana
o Chloroplasts also contain stroma, a dense fluid
✓ Visible light consists of wavelengths (including PHOTOSYNTHETIC PIGMENTS: THE LIGHT
those that drive photosynthesis) that produce RECEPTORS
colors we can ses
✓ PIGMENTS are subtances that absorb visible
light
✓ Different pigments absorb different
wavelengths
✓ Wavelengths that are not absorbed are reflected
or transmitted

CHLOROPLAST

✓ Chloroplast is surrounded by a double

membrane system.
✓ Inside the chloroplast are amorphous and gel-
like materials called the stroma.
✓ Embedded throughout the stroma are pigment
LEAF STRUCTURE containig thylakoids.
✓ Most leaves are efficient interceptors of light
because of their flat shape.

SPECTROPHOTOMETER

✓ Measures a pigment’s ability to absorb various

✓ Mesophyll cells containing chloroplasts and wavelengths
numerous intercellular spaces allows free ✓ Chlorophyll a is the main photosynthetic
diffusion and interchange of gasses atmosphere pigment
via epidermal openings called stomata.

Accesory pigments:

✓ Chlorophyll b, broaden the spectrum used for

photosynthesis
✓ Caratenoids absorb excessive light that would TWO TYPES OF PHOTOSYSTEM
damage chlorophyll
✓ Absorption spectrum is a graph plotting a 1. Photosystem II (PS II) functions first and is
pigment’s light absorption versus wavelength best. at absorbing a wavelength of 680 nm
✓ The absorption spectrum of chloropyhll a ➢ chlorophyll a reaction-center of PS II
suggests the violet-blue and red light work best is called P680
for photosynthesis 2. Photosystem I (PS I) is best at absorbing a
✓ Action spectrum profiles the relative wavelength of 700 nm
effectiveness of different wavelenghts of ➢ chlorophyll a reaction-center of PS I
radiation in driving a process is called P700

LINEAR/NON-CYCLIC ELECTRON FLOW

During the light reactions, there are two possible routes

for electron flow:

1. cyclic
2. linear

Linear electron flow, the primary pathway, involves both

PHOTOSYSTEM
photosystems (I & II) and produces ATP and NADPH
➢ Consists of a reaction-center complex (a type using light energy
of protein complex) surrounded by light
➢ Proteins in the thylakoid membrane
harvesting complexes
consists of 60 principal polypeptide
➢ Light-harvesting complexes (pigment
➢ 4 Supramolecular Transmembrane
molecules bound to proteins) funnel the energy
Complex
of photons to the reaction center
1. PS I
➢ A primary electron acceptor in the reaction
2. PS II
center accepts an excited electron from
3. Cyrochrome b/f complex
chlorophyll a
4. ATP synthase
➢ Solar-powered transfer of an electron from a
chlorophyll a molecule to the primary electron
acceptor is the first steps of the light reactions
1. A photon hits a pigment and its energy is passed
among pigment molecules until it excites P680
2. An excited electron from P680 is transferred to
the primary electron acceptor (Pheophytin)
3. P680+ is a very strong oxidizing agent
o H2O is split by enzymes, and the
electrins are transferred from the
hydrogen atoms to P680+, thus
reducing it to P680
o O2 is release as by-product of this
reaction
4. Each electron "falls" down an electron transport
chain from the primary electron acceptor of PS
II to PSI.
 o Energy released by the fall drives the intermembrane space and drive ATP synthesis
creation of a proton gradient across as they diffuse back into the mitochondrial
the thylakoid membrane matrix
5. Diffusion of H+ (protons) across the membrane ➢ In chloroplasts, protons are pumped into the
drives ATP synthesis thylakoid space and drive ATP synthesis as they
diffuse back into the stromainto the chemical
ELECTRON TRANSPORT CHAIN (ETC) energy of ATP
➢ Consist of a of series electron carrier molecules
and couples this electron transfer with the
transfer of protons (H+ ions) across a
membrane.

6. In PSI (like PS II), transferred light energy

excites P700, which loses an electron to an
electron acceptor
o P700+ accepts an electron passed
down from PS II via the electron
transport chain

CHEMIOSMOSIS

Energy of electron movement through the thylakoid

membrane creates and H+ gradient that drives ATP
sythesis.

7. Each electron "falls" down an electron transport chain

from the primary electron acceptor of PS I to the
protein ferredoxin (Fd)
8. The electrons are then transferred to NAP+ and reduce
it to NADPH

A COMPARISON OF CHEMIOSMOSIS IN
CHLOROPLASTS AND MITOCHONDRIA

➢ Chloroplasts and mitochondria generate ATP

chemiosmosis, but use different sources of
energy
➢ Mitochondria transfer chemical energy from
food to ATP; chloroplasts transform light
energy
➢ In mitochondria, protons are pumped to the
 CYCLIC ELECTRON FLOW

➢ Uses only photosystem I and produces ATP, but not

NADPH
➢ Generates surplus ATP, satisfying the higher demand
in the Calvin cycle
➢ Some organisms such as purple sulfur bacteria
have PS I but not PS II

PHOTOREGULATION AND ADAPTATION TO LIGHT

Photoinhibition

o involves damage of Reaction Center

complexes

Photooxidation

o involves Light Harvesting Center directly

Optimal use of light can be achieved via morphological,

structural, physiological, and biochemical characteristics such as:
C3 OR CALVIN CYCLE
➢ Pubescent leaves absorbs 40% less light.
➢ Chloroplast movement to the sides of the cells. ➢ CO2 is first incorporated into a 3-carbon compound.
➢ Morphogenetic adaptation of the leaf to light ➢ Stomata are open during the day.
conditions during growth ➢ Adaptive Value: more efficient than C4 and CAM
➢ Biochemical induction under light stress of enzymes plants under cool and moist conditions and under
Superoxide dismutase (SOD). normal light because requires less specialized
anatomy machinery (fewer enzymes and no
FROM LIGHT REACTION TO CALVIN CYCLE
specialized anatomy).
➢ CO2 Assimilation phase/ Photosynthetic Reaction ➢ Most plants are C3
(PCR) Cycle
ALTERNATIVE MECHANISMS OF CARBON
➢ Occurs in the stroma of the chloroplast
FIXATION HAVE EVOLVED IN HOT, ARID CLIMATES
➢ Entails the use of ATP (source chemical energy) &
NAPDH (source of reducing power) to reduce CO2 ➢ Dehydration is a problem of plants, sometimes
yielding glucose and other carbohydrates. requiring trade-offs with other metabolic processes,
especially photosynthesis.
PCR CYCLE
➢ On hot, dry days, plants close stomata, which
Three biochemical variants for the assimilation of CO2 conserves H2O but also limits photosythesis
➢ The closing of stomata reduces access to CO2 and
1. C3 or Calvin Cycle or reductive pentose pathway causes O2 to build up
2. C4 cycle or Hatch and Slack pathway ➢ These conditions favor a seemingly wasteful process
3. Crassulacean Acid Metabolism (CAM) pathway called photorespiration

CALVIN CYCLE PHOTORESPIRATION

• Carbon enters the cycle as CO2 and leaves as a sugar • In most plants (C3 plants), initial fixation of CO2, via
named glyceraldehyde-3-phospate (G3P) rubisco, forms three-carbon a compound
• For net synthesis of 1 G3P, the cycle must take place • In photorespiration, rubisco adds O2 instead of CO2
three times, fixing 3 molecules of CO2 in the Calvin cycle
• The Calvin cycle has three phases: • Photorespiration consumes O2 and organic fuel and
1. Carbon fixation (catalyzed by rubisco) releases CO2 without producing ATP or sugar
2. Reduction • Photorespiration or C2 photorespiratory carbon
3. Regeneration of the CO2 acceptor (RuBP) oxidation cycle (PCO) involves the cooperative
interaction of choloroplasts, peroxisiomes, and
mitochondria.
 C4 PLANTS ADAPTIVE VALUE

➢ Photosynthesizes faster than C3 plants under high

light intensity and high temperatures
➢ Has better Water Use Efficiency because PEP
Carboxylase brings in CO₂ faster and so does not
need to keep stomata open (less water lost by
transpiration);
➢ Example: occurs in grasses like Sugar cane, corn, and
many of our summer annual plants.

CRASSULACEAN ACID METABOLISM (CAM)

➢ Stomata open at night & Dark: Stomata opened

closed during the day
➢ The CO2 is converted to an acid and stored during the
C4 PLANTS OR HATCH AND SLACK PATHWAY night. During the day, the acid is broken down and the
CO2 is released to RUBISCO for photosynthesis
➢ CO₂ is first incorporated ➢ Cells have larger vacuoles, deeper stomates, and
into a compound. 4-carbon denser guard cells.
➢ Stomata are open during the day. ➢ Better water use efficiency tha C3 plants under arid
➢ Kranz Anatomy, distinct concentric arrangements of conditions due to opening stomata at night when
chloroplast containing bundle sheath cells transpiration rates are lower.
surrounding the vascular bundle ➢ Many succelents such as cactuses and agaves and also
some orchids and bromeliads.

➢ C4 plants minimize the cost of photorespiration by

incorporating CO2 into four carbon compounds in FACTORS AFFECTING PHOTOSYNTHESIS
mesophyll cells
➢ This step requires the enzyme PEP carbocylase The following may belimiting:
➢ PEP carboxylase has a highe affinity for CO2 than
rubisco does; it can fix CO2 even when CO2 • Light intensity
concentrataions are low • Carbon dioxide concentration
➢ These four-carbon compounds are exported to bundle- • Temperature
sheath cells, where they release CO2 that is then used • Water supply
in the Calvin Cycle • Chlorophyll concentration
• Enzyme concentration