Photosynthesis

David Lea-Smith
Lecturer of Microbiology
School of Biological Sciences
d.lea-smith@uea.ac.uk
 Outline
• What is photosynthesis?

• Advantages of photosynthesis

• Proteorhodopsin (simplest form of photosynthesis)

• Purple non-sulphur and sulphur bacteria

• Green sulphur bacteria

• Cyanobacteria, algae (and plants)

• Carbon fixation and light harvesting

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• Photoprotection
 What is photosynthesis?

Photosynthesis is a process used by plants and other

organisms to convert light energy into chemical energy that
can later be released to fuel the organisms' activities

Nearly every organism on the planet (including humans)

directly or indirectly relies on photosynthesis for energy
 The advantages of using sunlight as an energy
source

~3.5–7.0 kWh/m² per day Average UK household usage: 12.7 kWh per day
 The Electromagnetic Spectrum

• Energy of a single photon (a quantum of light) is greater in the UV end

of the spectrum than in the Infrared

• Shorter wavelength = higher frequency = higher energy

 The Electromagnetic Spectrum

• Energy of a single photon (a quantum of light) is greater in the UV end

of the spectrum than in the Infrared

• Shorter wavelength = higher frequency = higher energy

 The three photosynthetic processes in bacteria
Electron independent ATP generation- produces ATP

Cyclic electron transport- produces ATP

Linear electron transport- produces ATP + NADPH

 Certain compounds can absorb light energy

• Includes carotenoids, chlorophylls and bilins

• Contain a chromophore, a part of the compound where

light energy can be absorbed, resulting in production of an
excited electron

• Responsible for the colour of certain compounds

 Generating energy via rhodopsin proteins

• Light driven proton pump

• Involved in ion pumping, sensory

transduction and as photosensory
pigments (George Wald- Nobel
Prize- 1967)

• Bacteriorhodopsin (archaea)

• Proteorhodopsin (bacteria, archaea, eukaryotic algae)

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2
 Generating energy via rhodopsin proteins
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• Almost free ATP!

• All species that express proteorhodopsin are photoheterotrophs

(utilise both light energy and organic compounds as a source of
energy)
 Rhodopsin expressing organisms are highly
abundant
• In a recent study over 1,500 different proteo/bacterio-rhodopsin genes
were identified in the North Pacific (including in SAR11) (Olson et al,
2018)

• Also found in freshwater environments

• Role in enhancing ATP production, growth under nutrient limitation or

stress conditions

• Most absorb blue/green light (between 500 – 650 nm)

Microbial rhodopsins absorb light at different
spectra

(Ernst et al, 2013)

Other photosynthetic organisms predominantly
absorb light using chlorophylls

Chlorophyll a Porphyrin ring

Hydrocarbon tail

Chlorophyll b

What is the chemical difference

between chlorophyll a and b?
 Purple non-sulphur bacteria
• Belong to the Rhodospirillaceae group (Alphaproteobacteria)

• Include Rhodobacter sphaeroides and Rhodopseudomonas palustris

(genetically manipulatable)

Rhodopseudomonas palustris
(producing electricity from
waste)

• Can grow aerobically in the dark by respiring on organic carbons

sources
 Purple sulphur bacteria
• Belong to the Chromatiaceae and Ectothiorhodospiraceae groups
(Gammaproteobacteria)

• Can tolerate high sulphur environments

• Strictly anaerobic (no oxygen for respiration)

• Must fix carbon dioxide to survive
Purple bacteria absorb infrared light via
bacteriochlorophylls

Absorption spectra of R.
LH2 sphaeroides (Papagiannakis
et al, 2002)
Light energy is transferred to a reaction centre to
generate an excited electron

LH1 + RC Energy transfer between

bacteriochlorophylls
 Purple bacteria generate ATP via anoxygenic
photosynthesis

Not generating new electrons!

Intracytoplasmic
vesicles
(R. sphaeroides)
Stolz, 2001

Electron transport chain diagram (Minteer et al, 2014)

A closer look at intracytoplasmic vesicles

Scheuring et al, 2014

AFM images
Purple bacteria use ATP to fix carbon (and some
fix nitrogen too)
• Most purple non-sulphur bacteria are heterotrophs

• Some are photoautotrophs (Utilise chemical energy to fix carbon

dioxide into sugars)

• Some species also fix nitrogen

(Require anaerobic conditions)
Carbon and nitrogen fixation require an electron
donor
• Electrons are consumed by the nitrogenase and NADPH (required for
carbon fixation

• Reduced sulphur compounds, i.e. hydrogen sulphide, can act as

electron donors but are not widespread in most environments
 Green sulphur bacteria harvest light via a large
complex termed the chlorosome
• Strict anaerobic bacteria mostly
found in low light environments
Chlorobiaceae
 Green sulphur bacteria have similar electron
transport pathways to purple sulphur bacteria
 Oxygenic photosynthesis

Plants

Algae Cyanobacteria
Cyanobacteria and algae massively impact
ecosystems around the world

Desert surface Antarctica

Freshwater ecosystems Oceans

Cyanobacterial physiology
Chloroplasts descend from an internalized (eaten)
cyanobacterium

A cyanobacterium (UCYN-A) ‘captured’ by an

alga (haptophyte)
Chloroplasts descend from an internalized (eaten)
cyanobacterium
Algal physiology
 Algae take on all shapes and forms

• Algae can be as evolutionary diverse from each other as humans are

from yeast
 Oxygenic photosynthesis- photosystem II
• Uses water as an electron donor (far more abundant than H2S)
Oxygenic photosynthesis- electron transport to
photosystem I and generation of NADPH

Z-scheme
(FNR) Ferredoxin
(Cyt b6f) Cytochrome b6f
NADP reductase
(PQ) Plastoquinone (Fd) Ferredoxin

(PQH2) Plastoquinol
(PC) Plastocyanin
Oxygenic photosynthesis- electron transport to
photosystem I and generation of NADPH

Z-scheme
(FNR) Ferredoxin
(Cyt b6f) Cytochrome b6f
NADP reductase
(PQ) Plastoquinone (Fd) Ferredoxin

(PQH2) Plastoquinol
(PC) Plastocyanin
Oxygenic photosynthesis- Carbon fixation
Oxygenic photosynthesis- photorespiration
The Calvin Cycle
 Oxygenic photosynthesis- Cyclic electron
transport
• No electron input is required (Therefore no oxygen is produced)

• Only ATP is produced

Cyanobacteria (and maybe some plants)

Algae and some plants

Oxygenic photosynthesis- Light harvesting in
cyanobacteria and some algae
Oxygenic photosynthesis- Light harvesting in
cyanobacteria and some algae
Cyanobacteria are not always blue green
 Oxygenic photosynthesis- Light harvesting in
other algae and plants

• Contain chlorophyll (broaden the area of light harvested, not the

spectra (why plants are green and cyanobacteria are blue-green)
Cyanobacteria and algae are each responsible for
approximately a quarter of global primary
production

Plants: ~50%

Algae: ~25% Cyanobacteria: ~25%

Oxygenic photosynthesis changed the Earth’s
atmosphere
 It also made life less habitable for oxygenic
photosynthetic organisms
• Oxygen + excited electrons = reactive oxygen molecules

• Reactive oxygen molecules kill cells

• Carotenoids dissipate excess energy as heat

and fluorescence, and scavenge reactive
oxygen species (and also transfer energy
back to chlorophylls)

• Certain proteins also act as electron sinks (transfer oxygen to

protons to generate water)

• Integrated at each point of the electron transport chain

Photosynthetic organisms from different groups
often inhabit the same environment
Winogradsky column

O2
Cyanobacteria/algae

SO4 Aerobic
Nonsulfur phototrophic
bacteria water

Purple sulfur bacteria Anaerobic

water

Green sulfur bacteria

H2S Clostridia, Sediment

desulfovibrio Solid
 Summary
• Harvesting light for energy is key to survival of most
organisms on Earth

• Rhodopsins pump protons leading to ATP production

• Purple and green photosynthetic bacteria cycle electrons

leading to ATP production and use sulphur compounds to
donate electrons for NADH or ammonia production

• Oxygenic photosynthetic organisms use water as an

electron donor

• Oxygenic photosynthetic organisms use multiple

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mechanisms to dissipate excess energy and reactive

oxygen species