CAMPBELL
BIOLOGY
© 2014 Pearson Education, Inc.
TENTH EDITION CAMPBELL
BIOLOGY
Reece • Urry • Cain • Wasserman •
Minorsky • Jackson TENTH EDITION
11
Photosynthetic
Processes
Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick 李澤民 海洋植物生理暨生質能研究室 海洋生物科技暨資源學系 國立中山大學 高雄 台灣The Process That Feeds the Biosphere
Photosynthesis is the process that converts solar
energy into chemical energy
Directly or indirectly, photosynthesis nourishes
© 2014 Pearson Education, Inc.
Autotrophs sustain themselves without eating anything derived from other organisms
Autotrophs are the producers of the biosphere,
producing organic molecules from CO2 and other
inorganic molecules
Almost all plants are photoautotrophs, using the
© 2014 Pearson Education, Inc.
Figure 11.1a
Other organisms also benefit from photosynthesis.
Photosynthesis occurs in plants, algae, certain other unicellular eukaryotes, and some
prokaryotes
These organisms feed not only themselves but
© 2014 Pearson Education, Inc. Figure 11.2 (a) Plants (b) Multicellular alga (c) Unicellular eukaryotes (d) Cyanobacteria
(e) Purple sulfur bacteria 40 μm 1 μ m 11 μm
Figure 11.2a
© 2014 Pearson Education, Inc.
Figure 11.2b
Figure 11.2c
(c) Unicellular eukaryotes
11
μ
© 2014 Pearson Education, Inc.
Figure 11.2d
(d) Cyanobacteria
40 μm
Figure 11.2e
1
μ
m
(e) Purple sulfur bacteria
© 2014 Pearson Education, Inc.
相對於autotrophs
Heterotrophs obtain their organic material from other organisms
Heterotrophs are the consumers of the biosphere
Almost all heterotrophs, including humans, depend
Earth’s supply of fossil fuels was formed from the remains of organisms that died hundreds of
millions of years ago
In a sense, fossil fuels represent stores of solar
© 2014 Pearson Education, Inc.
Concept 11.1: Photosynthesis converts light
energy to the chemical energy of food
Chloroplasts are structurally similar to and likely
evolved from photosynthetic bacteria 藍綠菌
The structural organization of these organelles
allows for the chemical reactions of
photosynthesis 基質 類囊體
© 2014 Pearson Education, Inc. Figure 11.4 Stroma Granum Thylakoid Thylakoid space Outer membrane Intermembrane space Inner membrane 20 μm Stomata Chloroplast Mesophyll cell 1 μm Mesophyll Chloroplasts Vein Leaf cross section
Chloroplasts: The Sites of Photosynthesis
in Plants
Leaves are the major locations of photosynthesis
Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
Each mesophyll cell contains 30–40 chloroplasts
CO2 enters and O2 exits the leaf through
© 2014 Pearson Education, Inc.
Figure 11.4a
Leaf cross section
Stomata Chloroplast Mesophyll Chloroplasts Vein Mesophyll cell 20 μm CO2 O2
Figure 11.4d
20 μm Mesophyll
© 2014 Pearson Education, Inc.
A chloroplast has an envelope of two membranes
surrounding a dense fluid called the stroma
Thylakoids are connected sacs in the chloroplast which compose a third membrane system
Thylakoids may be stacked in columns called
grana
Chlorophyll, the pigment which gives leaves their
Figure 11.4b Chloroplast Stroma Granum Thylakoid Thylakoid space Outer membrane Intermembrane space Inner membrane 1 μm
© 2014 Pearson Education, Inc.
Figure 11.4c
Stroma Granum
Tracking Atoms Through Photosynthesis:
Scientific Inquiry
Photosynthesis is a complex series of reactions
that can be summarized as the following equation:
6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 + 6 H2O
The overall chemical change during
photosynthesis is the reverse of the one that occurs during cellular respiration
© 2014 Pearson Education, Inc.
The Splitting of Water
Chloroplasts split H2O into hydrogen and oxygen,
incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product
Figure 11.5
Reactants:
Products:
6 CO2 12 H2O
© 2014 Pearson Education, Inc.
Photosynthesis as a
Redox Process
Photosynthesis reverses the direction of electron
flow compared to respiration
Photosynthesis is a redox process in which H2O is
oxidized and CO2 is reduced
Photosynthesis is an endergonic process; the
Figure 11.UN01
becomes reduced
© 2014 Pearson Education, Inc.
The Two Stages of Photosynthesis: A Preview
Photosynthesis consists of the light reactions
(the photo part) and Calvin cycle (the synthesis
part)
The light reactions (in the thylakoids)
Split H2O
Release O2
Reduce the electron acceptor NADP+ to NADPH
The Calvin cycle (in the stroma) forms sugar from
CO2, using ATP and NADPH
The Calvin cycle begins with carbon fixation,
© 2014 Pearson Education, Inc. Figure 11.6-1 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O
Figure 11.6-2 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O NADPH ATP O2
© 2014 Pearson Education, Inc. Figure 11.6-3 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O O2 CO2 NADPH ATP CALVIN CYCLE
Figure 11.6-4 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O [CH2O] (sugar) CALVIN CYCLE CO2 NADPH ATP O2
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Concept 11.2: The light reactions convert solar
energy to the chemical energy of ATP and
NADPH
Chloroplasts are solar-powered chemical factories
Their thylakoids transform light energy into the chemical energy of ATP and NADPH
The Nature of Sunlight
Light is a form of electromagnetic energy, also
called electromagnetic radiation
Like other electromagnetic energy, light travels in
rhythmic waves
Wavelength is the distance between crests
of waves
Wavelength determines the type of
© 2014 Pearson Education, Inc.
The electromagnetic spectrum is the entire
range of electromagnetic energy, or radiation
Visible light consists of wavelengths (including those that drive photosynthesis) that produce colors we can see
Light also behaves as though it consists of
Figure 11.7
Visible light
Gamma
rays X-rays UV Infrared
Micro-waves Radio waves 380 450 500 550 600 650 700 750 nm Shorter wavelength
Higher energy Lower energy
Longer wavelength
11− 5 nm 11− 3 nm 1 nm 113 nm 116 nm (11 nm)9
1 m
© 2014 Pearson Education, Inc.
Photosynthetic
Pigments
: The Light Receptors
Pigments are substances that absorb visible light
Different pigments absorb different wavelengths
Wavelengths that are not absorbed are reflected
or transmitted
Leaves appear green because chlorophyll reflects
and transmits green light
Figure 11.8 Light Chloroplast Reflected light Granum Transmitted light Absorbed light
© 2014 Pearson Education, Inc.
A spectrophotometer measures a pigment’s ability to absorb various wavelengths
This machine sends light through pigments and
measures the fraction of light transmitted at each wavelength
© 2014 Pearson Education, Inc. Figure 11.9 White light Refracting prism Chlorophyll solution Photoelectric tube Galvanometer
The high transmittance (low
absorption) reading indicates that chlorophyll absorbs very
little green light. Slit moves to pass light
of selected wavelength.
Green light
Blue light
The low transmittance (high absorption) reading indicates that chlorophyll absorbs
most blue light.
1
2 3
Figure 11.10 Chloro-phyll a Chlorophyll b Carotenoids 400 500 600 700 Wavelength of light (nm) (a) Absorption spectra
A bs or pt ion of l ight by c hl or opl a s t pi gme nts Rat e o f p hotos y n the s is (me a s ur e d by O 2 release) 400 500 600 700 400 500 600 700 (b) Action spectrum Aerobic bacteria Filament of alga
© 2014 Pearson Education, Inc.
An absorption spectrum is a graph plotting a pigment’s light absorption versus wavelength
The absorption spectrum of chlorophyll a
suggests that violet-blue and red light work best
for photosynthesis
An action spectrum profiles the relative
effectiveness of different wavelengths of radiation in driving a process
Figure 11.10a Chloro-phyll a Chlorophyll b Carotenoids 400 500 600 700 Wavelength of light (nm) (a) Absorption spectra
A bsor pt ion of l ight by c hl or opl ast pi gm ent s
© 2014 Pearson Education, Inc. Figure 11.10b R at e of p hot osy nt hesi s (m easur ed by O 2 rel ease) 400 500 600 700 (b) Action spectrum
The action spectrum of photosynthesis was first demonstrated in 1883 by Theodor W. Engelmann
In his experiment, he exposed different segments
of a filamentous alga to different wavelengths
Areas receiving wavelengths favorable to
photosynthesis produced excess O2
He used the growth of aerobic bacteria clustered
© 2014 Pearson Education, Inc. Figure 11.10c 400 500 600 700 (c) Engelmann’s experiment Aerobic bacteria Filament of alga
Why so many chlorophyll pigments?
In response to different wavelength in
the environment!
1. shading
2. under forest
Chl b/Chl a increases (Chl b content
increases)
© 2014 Pearson Education, Inc.
Chlorophyll a is the main photosynthetic pigment
Chlorophyll b, broaden the spectrum used for photosynthesis
The difference in the absorption spectrum
between chlorophyll a and b is due to a slight
structural difference between the pigment molecules
Accessory pigments called carotenoids
absorb excessive light that would damage chlorophyll
Figure 11.11 Porphyrin ring: light-absorbing “head” of molecule; note magnesium atom at center Hydrocarbon tail:
interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown CH in chlorophyll a in chlorophyll b 3 CHO CH3
© 2014 Pearson Education, Inc.
Accessory pigments called carotenoids function in photoprotection; they absorb excessive light that would damage chlorophyll
Sun Radiation
Chlorophyll is the main photosynthetic pigment
Photosynthetically active radiation, often
abbreviated PAR,
designates the spectral range (wave band) of solar radiation from 400 to 700 nanometers that photosynthetic
organisms are able to use in the process of photosynthesis.
Phycobiliprotein: 藻膽蛋白
藍綠藻
紅藻
藻紅素
Phycoerthyrin
藻藍素
Phycocyanin
異藻藍素
Allophycocyanin
Phycobilisome藻膽蛋白體 in cyanobacteria
These antennae (called "phycobilisomes" in Synechococcus) are composed of pigment-proteins complexes arranged in such a way to capture light with a high
efficiency. Pigments that are bound to antenna systems may have very different colours (such as green, blue, pink or orange) and this will determine the wavelengths of the solar spectrum that cells can efficiently harvest in the oceanic waters.
complementary chromatic adaptation (CCA)
The Journal of Biological Chemistry, 276, 11449-11452 (2001)
Phycobiliproteins, bilin variation, and group III CA regulation.
Kehoe D M PNAS 2010;107:9029-9030
complementary chromatic
adaptation
Structure of a hemidiscoidal
phycobilisome of Tolypothrix tenuis
under different light conditions. (a) When illuminated by white light, the
phycobilisome contains phycoerythrin, phycocyanin, and allophycocyanin. Energy absorbed by phycoerythrin is transferred to phycocyanin and
allophycocyanin. The allophycocyanin core proteins are attached, via a linker protein, to the photosynthetic membrane, which is not shown. (b) When
illuminated by red light, the phycobilisome undergoes
complementary chromatic adaptation, in which phycoerythrin is no longer
produced but additional phycocyanin is produced. (After R. MacColl and D.
Guard-Friar, Phycobiliproteins, CRC illuminated by white light illuminated by red light
© 2014 Pearson Education, Inc.
Excitation of Chlorophyll by Light
When a pigment absorbs light, it goes from a
ground state to an excited state, which is unstable
When excited electrons fall back to the ground
state, photons are given off, an afterglow called
fluorescence
If illuminated, an isolated solution of chlorophyll
Figure 11.12
Excited state
Heat
(a) Excitation of isolated chlorophyll molecule (b) Fluorescence Ground state Photon (fluorescence) Photon Chlorophyll molecule E n er g y o f el ect ro n e−
© 2014 Pearson Education, Inc.
Figure 11.12a
A Photosystem: A Reaction-Center Complex
Associated with Light-Harvesting Complexes
A photosystem consists of a reaction-center
complex (a type of protein complex) surrounded by light-harvesting complexes
The light-harvesting complexes=Lhc (pigment
molecules bound to proteins) transfer the energy of photons to the reaction center
© 2014 Pearson Education, Inc.
Figure 11.13
(a) How a photosystem harvests light (b) Structure of a photosystem
Chlorophyll STROMA THYLA-KOID SPACE Protein subunits Thy la k oi d m e m bra ne Pigment molecules Primary electron acceptor Reaction-center complex STROMA Photosystem Light-harvesting complexes Photon Transfer
of energy Special pair of chloro-phyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) Thy la k oi d m e m bra ne e−
Figure 11.13a
(a) How a photosystem harvests light
Pigment molecules Primary electron acceptor Reaction-center complex STROMA Photosystem Light-harvesting complexes Photon Transfer
of energy Special pair of chloro-phyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) T h y lako id m em b ran e e−
© 2014 Pearson Education, Inc. Figure 11.13b (b) Structure of a photosystem Chlorophyll STROMA THYLA-KOID SPACE Protein subunits T h y lako id m em b ran e
Photosystem II (PSII)
Photosystem II contains chlorophylls a and b and absorbs light at
680nm. This is a large protein complex that is located in the thylakoid membrane.
LHC-II
MOST ABUNDANT MEMBRANE PROTEIN IN CHLOROPLASTS OF GREEN PLANTS
A TRANSMEMBRANE PROTEIN
BINDS
~ 7 CHLOROPHYLL a MOLECULES
~ 5 CHLOROPHYLL b MOLECULES
TWO CAROTENOIDS
COMPRISES ABOUT 50% OF ALL CHLOROPHYLL IN BIOSPHERE
© 2014 Pearson Education, Inc.
A primary electron acceptor in the reaction
center accepts excited electrons and is reduced as a result
Solar-powered transfer of an electron from a
chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions
There are two types of photosystems in the thylakoid membrane
Photosystem II (PS II) functions first (the
numbers reflect order of discovery) and is best at absorbing a wavelength of 680 nm
The reaction-center chlorophyll a of PS II is called
© 2014 Pearson Education, Inc.
Photosystem I (PS I) is best at absorbing a
wavelength of 700 nm
The reaction-center chlorophyll a of PS I is called
Linear Electron Flow
During the light reactions, there are two possible
routes for electron flow: cyclic and linear
Linear electron flow, the primary pathway,
involves both photosystems and produces ATP and NADPH using light energy
© 2014 Pearson Education, Inc.
There are 8 steps in linear electron flow:
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 (we now call it P680+)
Figure 11.UN02 Light H2O CO2 O2 LIGHT REACTIONS CALVIN CYCLE ATP NADPH ADP NADP+ [CH2O] (sugar)
© 2014 Pearson Education, Inc. Figure 11.14-1 Pigment molecules e− 1 2 P680 Light Photosystem II (PS II) Primary acceptor
Figure 11.14-2 Pigment molecules e− 1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 e− e− 2 H+ + O2 H2O ½
© 2014 Pearson Education, Inc. Figure 11.14-3 Pigment molecules e− 1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 e− e− 2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc ½
Figure 11.14-4 Pigment molecules e− 1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 e− e− 2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc P700 Light Photosystem I (PS I) 6 Primary acceptor e− ½
© 2014 Pearson Education, Inc. Figure 11.14-5 Pigment molecules e− 1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 e− e− 2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc P700 Light Photosystem I (PS I) 6 Primary acceptor e− e− 7 8 Fd e− Electron transport chain NADP+ reductase NADPH NADP+ + H+ ½
© 2014 Pearson Education, Inc.
3. H2O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680+, thus reducing it to P680
P680+ is the strongest known biological oxidizing agent
4. Each electron “falls” down an electron transport chain from the primary electron acceptor of PS II to PS I
5. Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane
Diffusion of H+ (protons) across the membrane drives ATP synthesis
© 2014 Pearson Education, Inc.
6. In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron
acceptor
P700+ (P700 that is missing an electron) accepts an electron passed down from PS II via the
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 NADP+ and reduce it to NADPH
The electrons of NADPH are available for the reactions of the Calvin cycle
© 2014 Pearson Education, Inc.
The energy changes of electrons during linear flow
through the light reactions can be shown in a mechanical analogy
Figure 11.15 Mill makes ATP NADPH Photosystem II Photosystem I ATP e− e− e− e− e− e− e−
© 2014 Pearson Education, Inc.
Cyclic Electron Flow
In cyclic electron flow, electrons cycle back from
Fd to the PS I reaction center
Cyclic electron flow uses only photosystem I and
produces ATP, but not NADPH
Figure 11.16 Primary acceptor Primary acceptor Fd Cytochrome complex Pc Pq Photosystem II Photosystem I Fd NADP+ + H+ NADP+ reductase NADPH ATP
© 2014 Pearson Education, Inc.
Some organisms such as purple sulfur bacteria
have PS I but not PS II
Cyclic electron flow is thought to have evolved
before linear electron flow
Cyclic electron flow may protect cells from
Cyclic Electron Flow
• Electron in Photosystem I is excited and transferred to ferredoxin that shuttles the electron to the
cytochrome complex.
A Comparison of
Chemiosmosis
in Chloroplasts
and Mitochondria
Chloroplasts and mitochondria generate ATP by
chemiosmosis, but use different sources of energy
Mitochondria transfer chemical energy from food
to ATP; chloroplasts transform light energy into the chemical energy of ATP
Spatial organization of chemiosmosis differs
between chloroplasts and mitochondria but also shows similarities
© 2014 Pearson Education, Inc.
In mitochondria, protons are pumped to the
intermembrane space and drive ATP synthesis as they diffuse back into the mitochondrial matrix
In chloroplasts, protons are pumped into the
thylakoid space and drive ATP synthesis as they diffuse back into the stroma
Figure 11.17 MITOCHONDRION STRUCTURE CHLOROPLAST STRUCTURE Thylakoid membrane Stroma ATP Thylakoid space Inter-membrane space Inner membrane Matrix Key Diffusion Electron transport chain ATP synthase ADP + H+ H+ Higher [H+] Lower [H+] Pi
© 2014 Pearson Education, Inc.
ATP and NADPH are produced on the side facing
the stroma, where the Calvin cycle takes place
In summary, light reactions generate ATP and
increase the potential energy of electrons by
Figure 11.18 Photosystem II Photosystem I Cytochrome complex Light Pq Light 4 H+ +2 H+ 4 H+ O2 H2O Pc Fd 3 2 1 NADP+ To Calvin Cycle NADP+ reductase STROMA (low H+ concentration) ATP synthase THYLAKOID SPACE (high H+ concentration) Thylakoid membrane ADP + H+ ATP Pi e−e − NADPH ½ + H+
© 2014 Pearson Education, Inc. Figure 11.18a STROMA (low H+ concentration) ATP ADP ATP synthase Pi + H+ THYLAKOID SPACE (high H+ concentration) 4 H+ Cytochrome complex Light Photosystem I Pc Pq 4 H+ Light Photosystem II 4 H+ Pc Fd Thylakoid membrane +2 H+ H2O O2 ½ e− e− 2 1
Figure 11.18b Cytochrome complex Light Photosystem I Pc Pq Fd NADP+ reductase NADP+ + H+ NADPH To Calvin Cycle STROMA (low H+ concentration) ATP ADP ATP synthase P + H+ THYLAKOID SPACE (high H+ concentration) 4 H+ 2 3
© 2014 Pearson Education, Inc.
Concept 11.3: The Calvin cycle uses the
chemical energy of ATP and NADPH to reduce
CO
2to sugar
The Calvin cycle, like the citric acid cycle,
regenerates its starting material after molecules enter and leave the cycle
The cycle builds sugar from smaller molecules by
using ATP and the reducing power of electrons carried by NADPH
Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P)
For net synthesis of 1 G3P, the cycle must take
place three times, fixing 3 molecules of CO2
The Calvin cycle has three phases
1. Carbon fixation (catalyzed by rubisco) 2. Reduction
© 2014 Pearson Education, Inc. Figure 11.UN03 Light H2O CO2 O2 LIGHT REACTIONS CALVIN CYCLE ATP NADPH ADP NADP+ [CH2O] (sugar)
Figure 11.19-1
Input 3 CO2, entering one per cycle Phase 1: Carbon fixation
Rubisco 3-Phosphoglycerate Calvin Cycle RuBP P P 3 P 3 P P 6
C3 cycle
© 2014 Pearson Education, Inc.
Figure 11.19-2
Input 3 CO2, entering one per cycle Phase 1: Carbon fixation
Rubisco 3-Phosphoglycerate 1,3-Bisphosphoglycerate Phase 2: Reduction G3P Calvin Cycle RuBP P P 3 P 3 P P 6 6 6 ADP P 6 P ATP P 6 6 6 NADP+ 6 Pi G3P P 1 Output Glucose and other organic compounds NADPH
Figure 11.19-3
Input 3 CO2, entering one per cycle Phase 1: Carbon fixation
Rubisco 3-Phosphoglycerate 1,3-Bisphosphoglycerate Phase 2: Reduction G3P Calvin Cycle G3P Phase 3: Regeneration of RuBP ATP 3 ADP 3 5 P RuBP P P 3 P 3 P P 6 6 6 ADP P 6 P ATP P 6 6 6 NADP+ 6 Pi G3P P 1 Glucose and other organic NADPH
© 2014 Pearson Education, Inc.
Concept 11.4: Alternative mechanisms of
carbon fixation have evolved in hot, arid
climates
Dehydration is a problem for plants, sometimes
requiring trade-offs with other metabolic processes, especially photosynthesis
On hot, dry days, plants close stomata, which
conserves H2O but also limits photosynthesis
The closing of stomata reduces access to CO2 and
causes O2 to build up
These conditions favor an apparently wasteful
Rubisco: Ribulose-1,5-bisphosphate
carboxylase/oxygenase
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Photorespiration: An Evolutionary Relic?
In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound
(3-phosphoglycerate)
In photorespiration, rubisco adds O2 instead of
CO2 in the Calvin cycle, producing a two-carbon
compound
Photorespiration consumes O2 and organic fuel
Photorespiration may be an evolutionary relic
because rubisco first evolved at a time when the
atmosphere had far less O2 and more CO2
Photorespiration limits damaging products of
light reactions that build up in the absence of the Calvin cycle
In many plants, photorespiration is a problem because on a hot, dry day it can drain as much as 50% of the carbon fixed by the Calvin cycle
© 2014 Pearson Education, Inc.
C
4Plants
C4 plants minimize the cost of photorespiration by
incorporating CO2 into four-carbon compounds
There are two distinct types of cells in the leaves
of C4 plants:
Bundle-sheath cells are arranged in tightly packed
sheaths around the veins of the leaf
Mesophyll cells are loosely packed between the bundle sheath and the leaf surface
Sugar production in C4 plants occurs in a three-step process:
1. The production of the four carbon precursors is catalyzed by the enzyme PEP carboxylase in the mesophyll cells
PEP carboxylase has a higher affinity for CO2 than rubisco does; it can fix CO2 even when CO2 concentrations are low
© 2014 Pearson Education, Inc.
2. These four-carbon compounds are exported to bundle-sheath cells
3. Within the bundle-sheath cells, they release CO2 that is then used in the Calvin cycle
Figure 11.20 Mesophyll cell Bundle-sheath cell Photo-synthetic cells of C4 plant leaf Vein (vascular tissue) C4 leaf anatomy Stoma The C4 pathway Mesophyll
cell PEP carboxylase
Oxaloacetate (4C) Malate (4C) Pyruvate (3C) CO2 ADP PEP (3C) ATP CO2 Calvin Cycle Bundle-sheath cell Sugar Vascular tissue
© 2014 Pearson Education, Inc. Figure 11.20a Mesophyll cell Bundle-sheath cell Photo-synthetic cells of C4 plant leaf Vein (vascular tissue) C4 leaf anatomy Stoma
Figure 11.20b
The C4 pathway
Mesophyll
cell PEP carboxylase
Oxaloacetate (4C) Malate (4C) Pyruvate (3C) CO2 ADP PEP (3C) ATP CO2 Calvin Cycle Bundle-sheath cell Sugar Vascular tissue
© 2014 Pearson Education, Inc.
Since the Industrial Revolution in the 1800s,
CO2 levels have risen greatly
Increasing levels of CO2 may affect C3 and C4
plants differently, perhaps changing the relative abundance of these species
The effects of such changes are unpredictable
CAM Plants
Some plants, including succulents, use
crassulacean acid metabolism (CAM) to fix carbon
CAM plants open their stomata at night,
incorporating CO2 into organic acids
Stomata close during the day, and CO2 is released
© 2014 Pearson Education, Inc. Figure 11.21 Sugarcane Pineapple C4 CO 2 CO2 CAM Organic acid Organic acid Night Day CO2 CO2 Calvin Cycle Calvin Cycle Sugar Sugar Bundle-sheath cell
(a) Spatial separation of steps (b) Temporal separation of steps Mesophyll cell 2 1 1 2
Figure 11.21a
© 2014 Pearson Education, Inc.
Figure 11.21b
The Importance of Photosynthesis: A Review
The energy entering chloroplasts as sunlight gets
stored as chemical energy in organic compounds
Sugar made in the chloroplasts supplies chemical
energy and carbon skeletons to synthesize the organic molecules of cells
Plants store excess sugar as starch in structures
such as roots, tubers, seeds, and fruits
In addition to food production, photosynthesis
© 2014 Pearson Education, Inc. Figure 11.22a O2 CO2 H2O Sucrose (export) H2O Light LIGHT REACTIONS: Photosystem II Electron transport chain Photosystem I Electron transport chain Chloroplast NADP+ ADP + Pi NADPH ATP RuBP G3P CALVIN CYCLE Starch (storage) 3-Phosphoglycerate Sucrose (export) O2 H2O Mesophyll cell CO2
Figure 11.22b
LIGHT REACTIONS CALVIN CYCLE REACTIONS
• Are carried out by molecules
in the thylakoid membranes
• Convert light energy to the
chemical energy of ATP and NADPH
• Split H2O and release O2 to the atmosphere
• Take place in the stroma
• Use ATP and NADPH to convert
CO2 to the sugar G3P
• Return ADP, inorganic phosphate,
© 2014 Pearson Education, Inc.
Figure 11.23
Flow of Genetic Information in the Cell:
DNA → RNA → Protein (Chapters 5–7)
Movement Across Cell Membranes (Chapter 7)
Energy Transformations in the Cell: Photosynthesis and Cellular
Respiration (Chapters 8–11) DNA mRNA Nucleus Nuclear pore Protein Ribosome mRNA Protein in vesicle Rough endoplasmic reticulum (ER) Vesicle forming Golgi apparatus Protein Plasma membrane Cell wall Photosynthesis in chloroplast Organic molecules Transport pump Cellular respiration in mitochondrion ATP ATP ATP ATP CO2 H2O H2O CO2 O2 O2 5 4 3 2 1 7 8 Vacuole 9 11 11 6 MAKE CONNECTIONS The Working Cell
Figure 11.23a
Flow of Genetic Information in the Cell: DNA → RNA → Protein (Chapters 5–7)
DNA mRNA Nucleus mRNA Nuclear pore Protein Protein in vesicle Rough endoplasmic reticulum (ER) Ribosome 1 2 3
© 2014 Pearson Education, Inc. Figure 11.23b Plasma membrane 4 Golgi apparatus Vesicle forming Protein Cell wall
Flow of Genetic Information in the Cell: DNA → RNA → Protein (Chapters 5–7)
5
Figure 11.23c Photosynthesis in chloroplast Organic molecules Cellular respiration in mitochondrion Transport pump Movement Across
Cell Membranes (Chapter 7) Energy Transformations in the Cell: Photosynthesis and Cellular Respiration (Chapters 8–11) O2 O2 H2O CO2 CO2 H2O ATP ATP ATP ATP 7 8 11 11 9
© 2014 Pearson Education, Inc.
Figure 11.UN04b
Corn plant surrounded by invasive velvetleaf plants
© 2014 Pearson Education, Inc. Figure 11.UN05 Primary acceptor Primary acceptor Pq Cytochrome complex Photosystem II Photosystem I Pc Fd NADPH NADP+ + H+ NADP+ reductase H2O O2
Figure 11.UN06 Regeneration of CO2 acceptor Calvin Cycle Carbon fixation Reduction 5 x 3C 3 x 5C 6 x 3C 3 CO2
© 2014 Pearson Education, Inc. Figure 11.UN07 pH 7 pH 4 pH 4 pH 8 ATP
Chlorophylls consist of a light-absorbing with a magnesium atom at the center and a long phytol tail that anchors the molecule in a membrane (Figure 1). They absorb light in the blue and red parts of the spectrum, but the green wavelengths are transmitted or reflected.
porphyrin ring