copyright © 2005 pearson education, inc. publishing as benjamin cummings november 27,...

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Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings November 27, 2012 Forgiveness Vindictive: revengeful; spiteful; showing a retaliatory spirit Do Now: Using only 2 sentence, respond to the statement below. Cut & paste the handout into your journal. The search for extrasolar planets has already detected rocky planets and interesting planetary candidates in the Habitable Zone of their host stars. Astrobiologists are pioneering new ways of imaging and examining these worlds, including how to assess their habitability. A new NAI-funded paper in the journal Astrobiology explores how a visible light filter photometry reading of an exoplanet might look if that planet had life on it. Such a reading of Earth, the pale blue dot, shows a “red edge” caused by green plants as they absorb light in the red part of the visible spectrum. But the characteristics of vegetation (or any organism with chlorophyll) on an exoplanet could vary depending on its host star. The signature of chlorophyll near a hot star could have a blue, rather than red, edge to protect a plant’s leaves from overheating. Or on a planet that orbits a cool, dim star chlorophyll may appear black as it tries to absorb as much light as possible across the whole range of the visible spectrum.

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Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

November 27, 2012 ForgivenessVindictive: revengeful; spiteful; showing a retaliatory spirit

Do Now: Using only 2 sentence, respond to the statement below. Cut & paste the handout into your journal.

The search for extrasolar planets has already detected rocky planets and interesting planetary candidates in the Habitable Zone of their host stars. Astrobiologists are pioneering new ways of imaging and examining these worlds, including how to assess their habitability. A new NAI-funded paper in the journal Astrobiology explores how a visible light filter photometry reading of an exoplanet might look if that planet had life on it. Such a reading of Earth, the pale blue dot, shows a “red edge” caused by green plants as they absorb light in the red part of the visible spectrum. But the characteristics of vegetation (or any organism with chlorophyll) on an exoplanet could vary depending on its host star. The signature of chlorophyll near a hot star could have a blue, rather than red, edge to protect a plant’s leaves from overheating. Or on a planet that orbits a cool, dim star chlorophyll may appear black as it tries to absorb as much light as possible across the whole range of the visible spectrum.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

PowerPoint Lectures for Biology, Seventh Edition

Neil Campbell and Jane Reece

Lectures by Chris Romero

Chapter 10Chapter 10

Photosynthesis

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Overview: The Process That Feeds the Biosphere

• Photosynthesis

– Is the process that converts solar energy into chemical energy

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Plants and other autotrophs

– Are the producers of the biosphere

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Plants are photoautotrophs

– They use the energy of sunlight to make organic molecules from water and carbon dioxide

Figure 10.1

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Photosynthesis

– Occurs in plants, algae, certain other protists, and some prokaryotes

These organisms use light energy to drive the synthesis of organic molecules from carbon dioxideand (in most cases) water. They feed not onlythemselves, but the entire living world. (a) Onland, plants are the predominant producers offood. In aquatic environments, photosyntheticorganisms include (b) multicellular algae, suchas this kelp; (c) some unicellular protists, suchas Euglena; (d) the prokaryotes calledcyanobacteria; and (e) other photosyntheticprokaryotes, such as these purple sulfurbacteria, which produce sulfur (sphericalglobules) (c, d, e: LMs).

(a) Plants

(b) Multicellular algae

(c) Unicellular protist 10 m

40 m(d) Cyanobacteria

1.5 m(e) Pruple sulfurbacteria

Figure 10.2

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Heterotrophs

– Obtain their organic material from other organisms

– Are the consumers of the biosphere

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

December 3, 2012 Caring

Terse: effectively brief and to the point; pithy; neatly concise; rudely short

1. Does dehydration synthesis (aka condensation synthesis) occur during the photosynthetic process?

2. How is energy stored within a plant?

3. What part does ATP play in photosynthesis?

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Concept 10.1: Photosynthesis converts light energy to the chemical energy of food

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Chloroplasts: The Sites of Photosynthesis in Plants

• The leaves of plants

– Are the major sites of photosynthesis

Vein

Leaf cross section

Figure 10.3

Mesophyll

CO2 O2Stomata

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Chloroplasts

– Are the organelles in which photosynthesis occurs

– Contain thylakoids and grana

Chloroplast

Mesophyll

5 µm

Outermembrane

Intermembranespace

Innermembrane

Thylakoidspace

ThylakoidGranumStroma

1 µm

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Tracking Atoms Through Photosynthesis: Scientific Inquiry

• Photosynthesis is summarized as

6 CO2 + 12 H2O + Light energy C6H12O6 + 6 O2 + 6 H2 O

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Splitting of Water

• Chloroplasts split water into

– Hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules

6 CO2 12 H2OReactants:

Products: C6H12O66 H2O 6 O2

Figure 10.4

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Photosynthesis as a Redox Process

• Photosynthesis is a redox process

– Water is oxidized, carbon dioxide is reduced

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Two Stages of Photosynthesis: A Preview

• Photosynthesis consists of two processes

– The light reactions

– The Calvin cycle

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The light reactions

– Occur in the grana

– Split water, release oxygen, produce ATP, and form NADPH

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The Calvin cycle

– Occurs in the stroma

– Forms sugar from carbon dioxide, using ATP for energy and NADPH for reducing power

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• An overview of photosynthesis

H2O CO2

Light

LIGHT REACTIONS

CALVINCYCLE

Chloroplast

[CH2O](sugar)

NADPH

NADP

ADP

+ P

O2Figure 10.5

ATP

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Concept 10.2: The light reactions convert solar energy to the chemical energy of ATP and NADPH

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Nature of Sunlight

• Light

– Is a form of electromagnetic energy, which travels in waves

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Wavelength

– Is the distance between the crests of waves

– Determines the type of electromagnetic energy

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The electromagnetic spectrum

– Is the entire range of electromagnetic energy, or radiation

Gammarays X-rays UV Infrared

Micro-waves

Radiowaves

10–5 nm 10–3 nm 1 nm 103 nm 106 nm1 m

106 nm 103 m

380 450 500 550 600 650 700 750 nm

Visible light

Shorter wavelength

Higher energy

Longer wavelength

Lower energyFigure 10.6

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The visible light spectrum

– Includes the colors of light we can see

– Includes the wavelengths that drive photosynthesis

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Photosynthetic Pigments: The Light Receptors

• Pigments

– Are substances that absorb visible light

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

– Reflect light, which include the colors we see

Light

ReflectedLight

Chloroplast

Absorbedlight

Granum

Transmittedlight

Figure 10.7

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The spectrophotometer

– Is a machine that sends light through pigments and measures the fraction of light transmitted at each wavelength

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• An absorption spectrum

– Is a graph plotting light absorption versus wavelength

Figure 10.8

Whitelight

Refractingprism

Chlorophyllsolution

Photoelectrictube

Galvanometer

Slit moves topass lightof selectedwavelength

Greenlight

The high transmittance(low absorption)reading indicates thatchlorophyll absorbsvery little green light.

The low transmittance(high absorption) readingchlorophyll absorbs most blue light.

Bluelight

1

2 3

40 100

0 100

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The absorption spectra of chloroplast pigments

– Provide clues to the relative effectiveness of different wavelengths for driving photosynthesis

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The absorption spectra of three types of pigments in chloroplasts Three different experiments helped reveal which wavelengths of light are photosynthetically important. The results are shown below.

EXPERIMENT

RESULTSA

bso

rptio

n o

f lig

ht

by

chlo

rop

last

pig

me

nts

Chlorophyll a

(a) Absorption spectra. The three curves show the wavelengths of light best absorbed by three types of chloroplast pigments.

Wavelength of light (nm)

Chlorophyll b

Carotenoids

Figure 10.9

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The action spectrum of a pigment

– Profiles the relative effectiveness of different wavelengths of radiation in driving photosynthesis

Rat

e o

f ph

otos

ynth

esis

(mea

sure

d by

O2 r

elea

se)

Action spectrum. This graph plots the rate of photosynthesis versus wavelength. The resulting action spectrum resembles the absorption spectrum for chlorophyll a but does not match exactly (see part a). This is partly due to the absorption of light by accessory pigments such as chlorophyll b and carotenoids.

(b)

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The action spectrum for photosynthesis

– Was first demonstrated by Theodor W. Engelmann

400 500 600 700

Aerobic bacteria

Filamentof alga

Engelmann‘s experiment. In 1883, Theodor W. Engelmann illuminated a filamentous alga with light that had been passed through a prism, exposing different segments of the alga to different wavelengths. He used aerobic bacteria, which concentrate near an oxygen source, to determine which segments of the alga were releasing the most O2 and thus photosynthesizing most.Bacteria congregated in greatest numbers around the parts of the alga illuminated with violet-blue or red light. Notice the close match of the bacterial distribution to the action spectrum in part b.

(c)

Light in the violet-blue and red portions of the spectrum are most effective in driving

photosynthesis.

CONCLUSION

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Chlorophyll a

– Is the main photosynthetic pigment

• Chlorophyll b

– Is an accessory pigment C

CH

CH2

CC

CC

C

CNNC

H3C

C

CC

C C

C

C

C

N

CC

C

C N

MgH

H3C

H

C CH2CH3

H

CH3C

HHCH2

CH2

CH2

H CH3

C O

O

O

O

O

CH3

CH3

CHO

in chlorophyll a

in chlorophyll b

Porphyrin ring:Light-absorbing“head” of moleculenote magnesiumatom at center

Hydrocarbon tail:interacts with hydrophobicregions of proteins insidethylakoid membranes ofchloroplasts: H atoms notshown

Figure 10.10

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Other accessory pigments

– Absorb different wavelengths of light and pass the energy to chlorophyll a

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Excitation of Chlorophyll by Light

• When a pigment absorbs light

– It goes from a ground state to an excited state, which is unstable

Excitedstate

Ene

rgy

of e

lect

ion

Heat

Photon(fluorescence)

Chlorophyllmolecule

GroundstatePhoton

e–

Figure 10.11 A

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• If an isolated solution of chlorophyll is illuminated

– It will fluoresce, giving off light and heat

Figure 10.11 B

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

A Photosystem: A Reaction Center Associated with Light-Harvesting Complexes

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• A photosystem

– Is composed of a reaction center surrounded by a number of light-harvesting complexes

Primary electionacceptor

Photon

Thylakoid

Light-harvestingcomplexes

Reactioncenter

Photosystem

STROMA

Thy

lako

id m

embr

ane

Transferof energy

Specialchlorophyll amolecules

Pigmentmolecules

THYLAKOID SPACE(INTERIOR OF THYLAKOID)Figure 10.12

e–

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The light-harvesting complexes

– Consist of pigment molecules bound to particular proteins

– Funnel the energy of photons of light to the reaction center

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• When a reaction-center chlorophyll molecule absorbs energy

– One of its electrons gets bumped up to a primary electron acceptor

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The thylakoid membrane

– Is populated by two types of photosystems, I and II

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Noncyclic Electron Flow

• Noncyclic electron flow

– Is the primary pathway of energy transformation in the light reactions

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Produces NADPH, ATP, and oxygen

Figure 10.13Photosystem II

(PS II)

Photosystem-I(PS I)

ATP

NADPH

NADP+

ADP

CALVINCYCLE

CO2H2O

O2 [CH2O] (sugar)

LIGHTREACTIONS

Light

Primaryacceptor

Pq

Cytochromecomplex

PC

e

P680

e–

e–

O2

+

H2O2 H+

Light

ATP

Primaryacceptor

Fd

ee–

NADP+

reductase

ElectronTransportchain

Electron transport chain

P700

Light

NADPH

NADP+

+ 2 H+

+ H+

1

5

7

2

3

4

6

8

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• A mechanical analogy for the light reactions

MillmakesATP

ATP

e–

e–e–

e–

e–

Pho

ton

Photosystem II Photosystem I

e–

e–

NADPH

Pho

ton

Figure 10.14 

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Cyclic Electron Flow

• Under certain conditions

– Photoexcited electrons take an alternative path

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• In cyclic electron flow

– Only photosystem I is used

– Only ATP is produced

Primaryacceptor

Pq

Fd

Cytochromecomplex

Pc

Primaryacceptor

Fd

NADP+

reductaseNADPH

ATPFigure 10.15

Photosystem II Photosystem I

NADP+

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria

• Chloroplasts and mitochondria

– Generate ATP by the same basic mechanism: chemiosmosis

– But use different sources of energy to accomplish this

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The spatial organization of chemiosmosis

– Differs in chloroplasts and mitochondria

Key

Higher [H+]Lower [H+]

Mitochondrion Chloroplast

MITOCHONDRIONSTRUCTURE

Intermembrancespace

Membrance

Matrix

Electrontransport

chain

H+ DiffusionThylakoidspace

Stroma

ATPH+

PADP+

ATPSynthase

CHLOROPLASTSTRUCTURE

Figure 10.16

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• In both organelles

– Redox reactions of electron transport chains generate a H+ gradient across a membrane

• ATP synthase

– Uses this proton-motive force to make ATP

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The light reactions and chemiosmosis: the organization of the thylakoid membrane

LIGHTREACTOR

NADP+

ADP

ATP

NADPH

CALVINCYCLE

[CH2O] (sugar)STROMA(Low H+ concentration)

Photosystem II

LIGHT

H2O CO2

Cytochromecomplex

O2

H2O O21

1⁄2

2

Photosystem ILight

THYLAKOID SPACE(High H+ concentration)

STROMA(Low H+ concentration)

Thylakoidmembrane

ATPsynthase

PqPc

Fd

NADP+

reductase

NADPH + H+

NADP+ + 2H+

ToCalvincycle

ADP

PATP

3

H+

2 H++2 H+

2 H+

Figure 10.17

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Concept 10.3: The Calvin cycle uses ATP and NADPH to convert CO2 to sugar

• The Calvin cycle

– Is similar to the citric acid cycle

– Occurs in the stroma

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The Calvin cycle has three phases

– Carbon fixation

– Reduction

– Regeneration of the CO2 acceptor

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The Calvin cycle

(G3P)

Input(Entering one

at a time)CO2

3

Rubisco

Short-livedintermediate

3 P P

3 P P

Ribulose bisphosphate(RuBP)

P

3-Phosphoglycerate

P6 P

6

1,3-Bisphoglycerate

6 NADPH

6 NADPH+

6 P

P6

Glyceraldehyde-3-phosphate(G3P)

6 ATP

3 ATP

3 ADP CALVINCYCLE

P5

P1

G3P(a sugar)Output

LightH2O CO2

LIGHTREACTION

ATP

NADPH

NADP+

ADP

[CH2O] (sugar)

CALVINCYCLE

Figure 10.18

O2

6 ADP

Glucose andother organiccompounds

Phase 1: Carbon fixation

Phase 2:Reduction

Phase 3:Regeneration ofthe CO2 acceptor(RuBP)

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Concept 10.4: Alternative mechanisms of carbon fixation have evolved in hot, arid climates

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• On hot, dry days, plants close their stomata

– Conserving water but limiting access to CO2

– Causing oxygen to build up

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Photorespiration: An Evolutionary Relic?

• In photorespiration

– O2 substitutes for CO2 in the active site of the enzyme rubisco

– The photosynthetic rate is reduced

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

C4 Plants

• C4 plants minimize the cost of photorespiration

– By incorporating CO2 into four carbon compounds in mesophyll cells

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• These four carbon compounds

– Are exported to bundle sheath cells, where they release CO2 used in the Calvin cycle

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• C4 leaf anatomy and the C4 pathway

CO2

Mesophyll cell

Bundle-sheathcell

Vein(vascular tissue)

Photosyntheticcells of C4 plantleaf

Stoma

Mesophyllcell

C4 leaf anatomy

PEP carboxylase

Oxaloacetate (4 C) PEP (3 C)

Malate (4 C)

ADP

ATP

Bundle-Sheathcell CO2

Pyruate (3 C)

CALVINCYCLE

Sugar

Vasculartissue

Figure 10.19

CO2

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

CAM Plants

• CAM plants

– Open their stomata at night, incorporating CO2 into organic acids

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• During the day, the stomata close

– And the CO2 is released from the organic acids for use in the Calvin cycle

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• The CAM pathway is similar to the C4 pathway

Spatial separation of steps. In C4 plants, carbon fixation and the Calvin cycle occur in differenttypes of cells.

(a) Temporal separation of steps. In CAM plants, carbon fixation and the Calvin cycle occur in the same cellsat different times.

(b)

PineappleSugarcane

Bundle-sheath cell

Mesophyll Cell

Organic acid

CALVINCYCLE

Sugar

CO2 CO2

Organic acid

CALVINCYCLE

Sugar

C4 CAM

CO2 incorporatedinto four-carbonorganic acids(carbon fixation)

Night

Day

1

2 Organic acidsrelease CO2 toCalvin cycle

Figure 10.20

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Importance of Photosynthesis: A Review

• A review of photosynthesis

Light 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

Calvin cycle reactions:• Take place in the stroma• Use ATP and NADPH to convert CO2 to the sugar G3P• Return ADP, inorganic phosphate, and NADP+ to the light reactions

O2

CO2H2O

Light

Light reaction Calvin cycle

NADP+

ADP

ATP

NADPH

+ P 1

RuBP 3-Phosphoglycerate

Amino acidsFatty acids

Starch(storage)

Sucrose (export)

G3P

Photosystem IIElectron transport chain

Photosystem I

Chloroplast

Figure 10.21

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Organic compounds produced by photosynthesis

– Provide the energy and building material for ecosystems