13 Photosynthesis

Key terms
absorption spectrum: the pattern of absorption of light of different wavelengths
accessory pigments: pigments other than chlorophyll present in a photosystem
action spectrum: the range of wavelengths of light within which a reaction occurs
ADP (adenosine diphosphate): a nucleotide, present in every living cell, made of adenosine and two
phosphate groups linked in series, and important in energy transfer reactions of metabolism
ATP (adenosine triphosphate): a nucleotide, present in every living cell, formed in photosynthesis and
respiration from ADP and Pi, and functioning in metabolism as a common intermediate between energy-
requiring and energy-yielding reactions
ATP synthase: (also called ATPase) an enzyme that catalyses the synthesis of ATP from ADP and Pi in
mitochondria (and chloroplasts)
autotrophic: capable of synthesising organic molecules from simple inorganic substances
C3 pathway: the light-independent stage of photosynthesis, in which carbon dioxide is fixed and the first
product is the three-carbon compound, glycerate 3-phosphate; C 3 plants fix carbon dioxide only by the
C3 pathway
C4 pathway: an additional pathway for carbon dioxide fixation that increases the supply of this raw material
of photosynthesis to the chloroplast; involves the formation of malate which then breaks down to produce
carbon dioxide; C4plants have this addition pathway for fixing carbon dioxide, they are often tropical plants
Calvin cycle: part of the light-independent stage of photosynthesis, it takes place in the stroma of
chloroplasts; it is the cycle of reactions in which ATP and reduced NADP provide energy and reducing
power to synthesise carbohydrate from carbon dioxide and in which the carbon dioxide acceptor molecule
(RuBP) is resynthesised
carotenoid: yellow or orange fat-soluble pigment, involved in photosynthesis as an accessory pigment
chemiosmosis: the process by which energy is transferred in the production of ATP, from the flow of
protons (H ) through ATP synthase
+

chlorophyll: the main photosynthetic pigment of green plants, occurs in the thylakoid membranes of the
grana in chloroplasts
chloroplast: organelle found in the cells of green plants, and in which the reactions of photosynthesis occur
chromatography: a technique for separating compounds in a mixture
competitive inhibitor: a substance which blocks or slows enzyme action by binding to the active site of the
enzyme
electron transport chain: a chain of adjacently placed protein carrier molecules in the thylakoid
membranes of the grana in the chloroplast (and in the inner mitochondrial membrane) along which electrons
pass by oxidation-reduction reactions; involved in the steps of photosynthesis in which light energy becomes
chemical energy
granum (plural: grana): stacked discs of thylakoid membranes found within the chloroplast, containing the
photosynthetic pigments; site of the light-dependent reaction of photosynthesis
light-dependent stage (of photosynthesis): reactions in which light energy is used split water and which
result in the formation of reduced NADP and ATP; occurs in the grana of chloroplasts
light-independent stage (of photosynthesis): reactions in which carbon dioxide is fixed and trioses are
formed using the products of the light-dependent stage; occurs in the stroma of chloroplasts
limiting factor: a factor which causes a biochemical process to proceed less rapidly than the maximum rate
that is possible, for example the concentration of carbon dioxide can be a limiting factor of photosynthesis
mesophyll: ground tissue cells containing chloroplasts
metabolism: integrated network of all the biochemical reactions of life
metabolite: a chemical substance involved in metabolism
NAD (nicotinamide adenine dinucleotide): a coenzyme involved in redox reactions, it acts as a hydrogen
acceptor; it is reduced in the link reaction and the Krebs cycle and then converted back to NAD in the
electron transport chain
palisade mesophyll: ground tissue cells of the leaf, containing chloroplasts; the site of photosynthesis
phosphate (Pi): phosphate ions, as involved in metabolism
photophosphorylation: the formation of ATP, using light energy; occurs in the grana of chloroplasts during
the light-dependent reactions of photosynthesis
photorespiration: a type of ‘wasteful’ respiration occurring in green plants in conditions of high light and
low carbon dioxide concentration
photosystem: a multimoleculer complex in the thylakoid membranes of chloroplasts, involved in the light-
dependent reactions of photosynthesis
reaction centre: a single molecule of chlorophyll a into which the other pigment molecules of a
photosystem funnel energy
respiration: the cellular process by which sugars and other substances are broken down to release chemical
energy for other cellular processes
ribulose bisphosphate: the five-carbon acceptor molecule for carbon dioxide in the light-independent stage
of photosynthesis
Rubisco (ribulose bisphosphate carboxylase): a five-carbon sugar which acts as a receptor molecule for
carbon dioxide in the light-independent reactions of photosynthesis
stroma: the interior of the chloroplast, between the grana, where the thylakoid membranes are few and are
loosely arranged; site of the light-independent reactions of photosynthesis
sugars: compounds with the general formula Cx(H2O)y, where x is approximately equal to y, and containing
an aldehyde or a ketone group
thylakoid: the membrane system of a chloroplast
 Topic summary
• Photosynthesis is an energy transfer process in which green plants
manufacture carbohydrates from carbon dioxide and water, using energy from sunlight. Oxygen is the
waste product. Chloroplasts, the organelles in which photosynthesis occurs, are chiefly found in the
mesophyll cells of the leaves, particularly the palisade mesophyll cells. Plants use the products of
photosynthesis to manufacture the other carbohydrates, lipids, proteins and all other metabolites, enzymes
and pigments required. For this they require energy transferred by respiration and selected mineral ions,
absorbed from the soil. Consequently, plant nutrition is described as autotrophic (self-feeding).
• The leaf is the ‘factory’ for photosynthesis, with palisade cells in positions to absorb maximum light. The
cells are supplied with water from the xylem vessels, and with carbon dioxide from the air by diffusion
through open stomata. The products of photosynthesis are exported from the leaf in the sieve tubes in the
phloem.
• Chloroplasts are enclosed by a double membrane. The inner membrane intucks to form the membrane
systems of the chloroplast (thylakoid membranes). These are arranged in compact, disc-shaped piles
called grana, around which are loosely arranged thylakoid membranes of the stroma.
• Chlorophyll consists of a mixture of pigments, two of which are chlorophylls. They can be separated
by chromatography. Measurements of the absorption spectrum of chlorophyll and the action
spectrum of photosynthesis show that it is the blue and red components of white light that are selectively
absorbed and are most effective in bringing about photosynthesis.
• Photosynthesis can be divided into two, linked stages.
• A light-dependent stage occurs in the grana and results in the formation of reduced NADP and ATP. It
involves the photolysis of water and the release of oxygen.
• A light-independent stage occurs in the stroma and involves the fixing of carbon dioxide to form trioses,
using the products of the light-dependent stage. Trioses are used in a variety of biochemical reactions within
the chloroplast to form monosaccharides (glucose and fructose) that are converted to sucrose for
translocation or to starch for storage. Other reactions use minerals from the soil to form amino acids that are
translocated and used for protein synthesis.
• In green plants the volume of oxygen-enriched gas given off in the light is a measure of the rate of
photosynthesis. The volume of gas produced by aquatic plants under different conditions can be measured
using a microburette (photosynthometer).
• As photosynthesis consists of a number of interconnected reactions, the slowest will determine the overall
rate. The factor limiting its rate is then described as the limiting factor. A limited supply of either carbon
dioxide or light will limit the overall rate of photosynthesis, since both are essential for photosynthesis.
• The autotrophic nutrition of the green plant also sustains other living things, since other organisms feed on
plants or plant products, directly or indirectly.
• Plants, including crop plants show adaptations to different environments. These include C4 carbon dioxide
metabolism as part of photosynthesis in many successful tropical and sub-tropical plants. This mechanism
is a response to the harmful effects of high oxygen concentrations and low carbon dioxide concentrations on
the carbon dioxide-fixing enzyme, rubisco, at high ambient temperatures. The structural adaptations shown
by mesophyll tissue in the leaves of sorghum and maize provide the venue for C4 carbon dioxide fixation.
 Data handling
Temperature and the rate of photosynthesis
The effect of temperature on the amount of gas given off by an aquatic plant shoot (stem and leaves) in the
light can be investigated, using the apparatus shown in Figure 10.17 (page 272).
A freshly cut shoot of pondweed is used which, when inverted, produces a vigorous stream of gas bubbles
from the base. The bubbles confirm the pondweed is actively photosynthesising. The apparatus contains a
dilute solution of sodium hydrogencarbonate, which supplies the carbon dioxide (as – HCO3 ions) required
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by the plant for photosynthesis. The quantity of oxygen evolved in a given time (30 minutes) is a measure of
how much photosynthesis has occurred in that period. This was estimated by collecting the gas bubbles
produced into the capillary tube and measuring the length of the resulting bubble. This length is then
converted to a volume.
In one experiment the effect of temperature on the rate of photosynthesis at both high and low light
intensities was measured. Light intensity was varied by having the light source at different distances from
the pond weed. The result of one experiment is given in the table below.
The effect of temperature on photosynthesis at high and low light intensities

Temperature Oxygen produced in

/ °C 30 minutes / cm
3

at high at low light

light intensity
intensity

5 26 20

10 39 25

15 66 28

17.5 95 30

20 128 33

22.5 110 30

25 76 28
a) Construct a graph to show the effects of temperature on gas production at high and low light intensities.
b) Describe in your own words the effect of the rise in temperature from 10°C to 20°C on gas production by
pond weed in high and low light intensities.
c) Suggest the likely cause of the change in the rate of photosynthesis at the higher temperatures.
d) The result of this experiment was interpreted as showing that photosynthesis is made up of two sequential
steps or reactions. Explain what these reactions are and why a rise in temperature has a different effect on
the two reactions.
 Do and understand
Experimental design
The diagram below shows a method of investigating photosynthesis in green plant leaf tissue.
Examine the steps carefully and then answer the following questions.
a) i) Explain why the leaf discs initially float in the hydrogencarbonate solution.
ii) Identify where the gas bubbles come from when the plunger is drawn back and the barrel contents are placed
under tension (negative pressure).
iii) Describe why the illuminated discs rise in the barrel after illumination.
b) Design an experiment to investigate the effect of different carbon dioxide concentrations on the rate of
photosynthesis, using this procedure. In your account pay particular attention to:
i) the safety steps that are necessary
ii) how you best assure yourself the results will be significant.
 Suggested website
• www.saddleworth.oldham.sch.uk/science/simulations/waterweed.htm – A simulation of the
photosynthometer can be used to investigate the effects of environmental factors (light intensity,
CO2 concentration and temperature) on the rate of photosynthesis. It generates results to present and discuss,
as an alternative exercise to a laboratory practical on photosynthesis.

Further reading
Articles in recent editions of Biological Sciences Review:
• ‘The plastid family’, 24.3, pp 20–25 (Feb 2012)
• ‘Photosynthesis’, 23.1, pp 20–21 (Sept 2010)
• ‘What is: A limiting factor?’, 24.2, pp 30–32 (Nov 2011)
• ‘Rubisco – the world’s most important enzyme’, 23.4, pp 2–5 (April 2011)