• 沒有找到結果。

光合作用

N/A
N/A
Protected

Academic year: 2021

Share "光合作用"

Copied!
148
0
0

加載中.... (立即查看全文)

全文

(1)

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

(2)

The Process That Feeds the Biosphere

Photosynthesis is the process that converts solar

energy into chemical energy

 Directly or indirectly, photosynthesis nourishes

(3)

© 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

(4)
(5)

© 2014 Pearson Education, Inc.

Figure 11.1a

Other organisms also benefit from photosynthesis.

(6)

 Photosynthesis occurs in plants, algae, certain other unicellular eukaryotes, and some

prokaryotes

 These organisms feed not only themselves but

(7)

© 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

(8)

Figure 11.2a

(9)

© 2014 Pearson Education, Inc.

Figure 11.2b

(10)

Figure 11.2c

(c) Unicellular eukaryotes

11

μ

(11)

© 2014 Pearson Education, Inc.

Figure 11.2d

(d) Cyanobacteria

40 μm

(12)

Figure 11.2e

1

μ

m

(e) Purple sulfur bacteria

(13)

© 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

(14)

 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

(15)

© 2014 Pearson Education, Inc.

(16)

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

(17)

© 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

(18)

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

(19)

© 2014 Pearson Education, Inc.

Figure 11.4a

Leaf cross section

Stomata Chloroplast Mesophyll Chloroplasts Vein Mesophyll cell 20 μm CO2 O2

(20)

Figure 11.4d

20 μm Mesophyll

(21)

© 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

(22)

Figure 11.4b Chloroplast Stroma Granum Thylakoid Thylakoid space Outer membrane Intermembrane space Inner membrane 1 μm

(23)

© 2014 Pearson Education, Inc.

Figure 11.4c

Stroma Granum

(24)

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

(25)

© 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

(26)

Figure 11.5

Reactants:

Products:

6 CO2 12 H2O

(27)

© 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

(28)

Figure 11.UN01

becomes reduced

(29)

© 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

(30)

 The Calvin cycle (in the stroma) forms sugar from

CO2, using ATP and NADPH

 The Calvin cycle begins with carbon fixation,

(31)

© 2014 Pearson Education, Inc. Figure 11.6-1 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O

(32)

Figure 11.6-2 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O NADPH ATP O2

(33)

© 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

(34)

Figure 11.6-4 Light Thylakoid Stroma Chloroplast LIGHT REACTIONS NADP+ ADP P i + H2O [CH2O] (sugar) CALVIN CYCLE CO2 NADPH ATP O2

(35)

© 2014 Pearson Education, Inc.

(36)
(37)

© 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

(38)

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

(39)

© 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

(40)

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

(41)

© 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

(42)

Figure 11.8 Light Chloroplast Reflected light Granum Transmitted light Absorbed light

(43)

© 2014 Pearson Education, Inc.

(44)

 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

(45)

© 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

(46)

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

(47)

© 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

(48)

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

(49)

© 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

(50)

 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

(51)

© 2014 Pearson Education, Inc. Figure 11.10c 400 500 600 700 (c) Engelmann’s experiment Aerobic bacteria Filament of alga

(52)

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)

(53)

© 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

(54)

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

(55)

© 2014 Pearson Education, Inc.

(56)

Accessory pigments called carotenoids function in photoprotection; they absorb excessive light that would damage chlorophyll

(57)

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.

(58)
(59)
(60)
(61)

Phycobiliprotein: 藻膽蛋白

藍綠藻

紅藻

藻紅素

Phycoerthyrin

藻藍素

Phycocyanin

異藻藍素

Allophycocyanin

(62)

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.

(63)
(64)

complementary chromatic adaptation (CCA)

The Journal of Biological Chemistry, 276, 11449-11452 (2001)

(65)

Phycobiliproteins, bilin variation, and group III CA regulation.

Kehoe D M PNAS 2010;107:9029-9030

(66)

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

(67)

© 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

(68)

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

(69)

© 2014 Pearson Education, Inc.

Figure 11.12a

(70)
(71)
(72)

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

(73)

© 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

(74)

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

(75)

© 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

(76)

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.

(77)

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

(78)
(79)
(80)
(81)

© 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

(82)

 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

(83)

© 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

(84)

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

(85)

© 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+)

(86)

Figure 11.UN02 Light H2O CO2 O2 LIGHT REACTIONS CALVIN CYCLE ATP NADPH ADP NADP+ [CH2O] (sugar)

(87)

© 2014 Pearson Education, Inc. Figure 11.14-1 Pigment molecules e1 2 P680 Light Photosystem II (PS II) Primary acceptor

(88)

Figure 11.14-2 Pigment molecules e1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 ee2 H+ + O2 H2O ½

(89)

© 2014 Pearson Education, Inc. Figure 11.14-3 Pigment molecules e1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 ee2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc ½

(90)

Figure 11.14-4 Pigment molecules e1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 ee2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc P700 Light Photosystem I (PS I) 6 Primary acceptor e½

(91)

© 2014 Pearson Education, Inc. Figure 11.14-5 Pigment molecules e1 2 P680 Light Photosystem II (PS II) Primary acceptor 3 ee2 H+ + O2 H2O ATP 4 5 Electron transport chain Cytochrome complex Pq Pc P700 Light Photosystem I (PS I) 6 Primary acceptor ee7 8 Fd eElectron transport chain NADP+ reductase NADPH NADP+ + H+ ½

(92)
(93)

© 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

(94)

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

(95)

© 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

(96)

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

(97)

© 2014 Pearson Education, Inc.

 The energy changes of electrons during linear flow

through the light reactions can be shown in a mechanical analogy

(98)

Figure 11.15 Mill makes ATP NADPH Photosystem II Photosystem I ATP eeeeeee

(99)

© 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

(100)

Figure 11.16 Primary acceptor Primary acceptor Fd Cytochrome complex Pc Pq Photosystem II Photosystem I Fd NADP+ + H+ NADP+ reductase NADPH ATP

(101)

© 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

(102)

Cyclic Electron Flow

• Electron in Photosystem I is excited and transferred to ferredoxin that shuttles the electron to the

cytochrome complex.

(103)
(104)

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

(105)

© 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

(106)

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

(107)

© 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

(108)

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 eeNADPH ½ + H+

(109)

© 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 ee2 1

(110)

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

(111)

© 2014 Pearson Education, Inc.

Concept 11.3: The Calvin cycle uses the

chemical energy of ATP and NADPH to reduce

CO

2

to 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

(112)

 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

(113)

© 2014 Pearson Education, Inc. Figure 11.UN03 Light H2O CO2 O2 LIGHT REACTIONS CALVIN CYCLE ATP NADPH ADP NADP+ [CH2O] (sugar)

(114)

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

(115)

© 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

(116)

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

(117)

© 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

(118)

Rubisco: Ribulose-1,5-bisphosphate

carboxylase/oxygenase

(119)

© 2014 Pearson Education, Inc.

(120)
(121)

© 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

(122)

 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

(123)

© 2014 Pearson Education, Inc.

C

4

Plants

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

(124)

 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

(125)

© 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

(126)

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

(127)

© 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

(128)

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

(129)

© 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

(130)

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

(131)

© 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

(132)

Figure 11.21a

(133)

© 2014 Pearson Education, Inc.

Figure 11.21b

(134)

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

(135)

© 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

(136)

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,

(137)

© 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

(138)

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

(139)

© 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

(140)

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

(141)

© 2014 Pearson Education, Inc.

(142)

Figure 11.UN04b

Corn plant surrounded by invasive velvetleaf plants

(143)

© 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

(144)

Figure 11.UN06 Regeneration of CO2 acceptor Calvin Cycle Carbon fixation Reduction 5 x 3C 3 x 5C 6 x 3C 3 CO2

(145)

© 2014 Pearson Education, Inc. Figure 11.UN07 pH 7 pH 4 pH 4 pH 8 ATP

(146)
(147)
(148)

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

參考文獻

相關文件

Figure 3: Case 3 description: (a) gross specimen; (b) scanning power view showing numerous endothelial cell lined proliferating capillaries and dense inflammatory cell infiltration

Figure 4: (a) Intraoperative photograph showing ligation of branches of external carotid artery, (b) intraoperative photograph during resection of carotid body paragangliomas

Regular follow-up, during 8 months, revealed clinical healing after 4 months with disappearance of mandibular swelling (Figure 5(a)), periapical lesion regression after 8 months

(A) The scene where the heroine of Guardian: The Lonely and Great God places a maple leaf between the pages of a book was shot in Canada.. (B) The maple leaf on the Canadian

Figure 1.5 Some consumer applications of computer vision: (a) image stitching: merging different views (Szeliski and Shum 1997) c 1997 ACM; (b) exposure bracketing: merging

Thus any continuous vector function r defines a space curve C that is traced out by the tip of the moving vector r(t), as shown in Figure 1.... The curve, shown in Figure 2,

 Construction and Evaluation of Actuarial Models (Exam

   茶捲葉蛾多危害成葉,幼蟲分散後隨即吐絲將 2 片葉黏在一起,棲於內面 取食,隨著幼蟲長大,再將附近