Biology The Dynamic Science 3rd Edition

Russell
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9
PHOTOSYNTHESIS

Chapter Outline
WHY IT MATTERS
9.1 PHOTOSYNTHESIS: AN OVERVIEW
Electrons play a primary role in photosynthesis.
Oxygen released by photosynthesis derives from the splitting of water.
In eukaryotes, photosynthesis takes place in chloroplasts.
9.2 THE LIGHT-DEPENDENT REACTIONS OF PHOTOSYNTHESIS
Electrons in pigment molecules absorb light energy in photosynthesis.
Chlorophylls are the main light receptor pigments in photosynthesis.
The photosynthetic pigments are organized into photosystems in chloroplasts.
Electron flow from water to NADP+ leads to the synthesis of NADPH and ATP.
Electron flow can also drive ATP synthesis by flowing cyclically around photosystem I.
Experiments with chloroplasts helped confirm the synthesis of ATP by chemiosmosis.
9.3 THE LIGHT-INDEPENDENT REACTIONS OF PHOTOSYNTHESIS
The Calvin cycle uses NADPH, ATP, and CO2 to generate carbohydrates.
G3P is the starting point for synthesis of many other organic molecules.
Rubisco is the key enzyme of the world’s food economy.
FOCUS ON BASIC RESEARCH: TWO-DIMENSIONAL PAPER CHROMATOGRAPHY AND THE CALVIN
CYCLE
INSIGHTS FORM THE MOLECULAR REVOLUTION: SMALL BUT PUSHY: WHAT IS THE FUNCTION OF
THE SMALL SUBUNIT OF RUBISCO?
9.4 PHOTORESPIRATION AND ALTERNATIVE PROCESSES OF CARBON FIXATION
Photorespiration produces carbon dioxide that is used by the Calvin cycle.
Alternative processes of carbon fixation minimize photorespiration.
9.5 PHOTOSYNTHESIS AND CELLULAR RESPIRATION COMPARED

82 Chapter Nine
 Photosynthesis and cellular respiration are similar processes with reactions that essentially are the reverse of
each other.
Evolution of photosynthesis and respiration.
UNANSWERED QUESTIONS

Learning Objectives
After reading the chapter, students should be able to:
Understand the basic properties of light and how sunlight specifically affects pigments in plants.
Define where photosynthesis takes place and the two stages of photosynthesis.
Explain the importance of the proton gradient and why a final electron acceptor is necessary to maintain the gradient.
Analyze the difference between the cyclic and noncyclic reactions of photosynthesis.
Understand the importance of the generation of NADPH and the utilization of ATP in the reactions of photosynthesis.
Know the steps of the light-dependent and light-independent reactions. Know the raw materials (reactants) needed to
start each step, the products made by each step, and where in the plant cell each step occurs.
Explain the difference between cyclic and noncyclic pathways in photosynthesis.
Explain how autotrophs use the intermediates as well as the products of photosynthesis in their own metabolism.
Apply the knowledge of photosynthesis to the impact autotrophs living in the oceans and on land have on global
processes such as carbon cycling. Understand how this activity influences the global climate and discuss the role
humans play in these events.
Explain the role of Rubisco to the world’s food economy.

Key Terms
photosynthesis electromagnetic reaction center Photorespiration
autotrophs spectrum linear electron flow C4 pathway
photoautotrophs chlorophylls photophosphorylation crassulacean acid
heterotrophs carotenoids cyclic electron flow metabolism

light-dependent absorption spectrum Calvin cycle CAM pathway

reactions action spectrum RuBP carboxylase,
light-independent photosystems oxygenase (rubisco)
reactions (Calvin photosystems II and I C3 pathway
cycle)
antenna complex C3 plants
CO2fixation

Lecture Outline
Why It Matters
A. Plants and algae use light energy to produce organic molecules, like sugar, by a process called
photosynthesis.
B. Englemann’s experiment (Figure 9.1) with photosynthesis of filamentous alga:
1. Violet, blue, and red light colors caused most oxygen to be released.
2. Bacteria clustered around the areas that released most oxygen.
C. Photosynthetic organisms include plants, some protists (algae), some archaeans, and some bacteria.
D. Most photosynthetic organisms use radiant energy of sunlight, convert to chemical energy, and form
complex organic chemicals.

9.1 Photosynthesis: An Overview

A. Plants are primary producers (autotrophs) that produce organic molecules and serve to provide food to
animals, the consumers (heterotrophs).

Photosynthesis 83
 B. The bodies of plants and animals eventually are a source of organic chemicals for the decomposers.
C. Electrons play a primary role in photosynthesis.
1. In light-dependent reactions, light energy is absorbed and converted into chemical energy: ATP and
NADPH.
2. In light-independent reactions (i.e., Calvin cycle):
a. Energy from ATP and electrons of NADPH are used for CO 2 fixation.
b. Added electrons and protons convert this into a sugar.
3. In plants, algae, and cyanobacteria, the source of electrons and protons for fixing CO 2 is water.
4. When water splits, oxygen is released as a by-product.
5. Photosynthesis primarily produces glucose, but other organic molecules are also assembled directly or
indirectly as products of photosynthesis.
D. Oxygen released by photosynthesis derives from the splitting of water.
1. Samuel Ruben and Martin Kamen of the University of California at Berkeley used a heavy isotope of
oxygen to trace the pathway of oxygen during photosynthesis.
a. Oxygen comes from the splitting of water, not the CO2.
2. The water-splitting reaction probably developed before oxygen-consuming organisms evolved, and led
to a significant change in the early Earth’s atmosphere.
E. In eukaryotes, photosynthesis takes place in chloroplasts.
1. Photosynthesis reactions take place in chloroplasts of plants and algae.
2. Photosynthesis reactions in bacteria are distributed on plasma membrane and cytosol.
3. Chloroplast.
a. Consist of an outer and an inner membrane covering the entire surface of the organelle.
b. Between these is an intermembrane compartment.
c. Fluid in the middle is called the stroma.
d. In the stroma are several flattened sacs called thylakoids with enclosed space, the thylakoid lumen.
e. Stack of thylakoids is called granum (Figure 9.3).
f. The grana are interconnected by tubular membranes called stromal lamellae.
g. The thylakoid membranes and stromal lamellae house the molecules that carry out the light-
dependent reactions, including the light reaction pigments, electron transfer carriers, and ATP
synthase enzymes.
h. Stomata bring CO2 into cells containing chloroplasts.
i. Water and minerals are transported through the roots.

9.2 The Light-Dependent Reactions of Photosynthesis

A. These involve two main processes: light absorption and synthesis of NADPH and ATP.
B. Electrons in pigment molecules absorb light energy in photosynthesis.
1. Visible light is a small part of the electromagnetic spectrum (Figure 9.4).
2. Shorter wavelengths have greater energy.
3. Humans see white light: a mixture of the entire visible spectrum ranging from 700nm to 400 nm.
4. Light energy travels in the form of a photon, which is a discrete unit that contains a fixed amount of
energy.
5. Energy in a photon is inversely proportional to the wavelength: shorter the wavelength, higher the
energy; longer the wavelength, lesser the energy.
6. Chlorophylls (yellow-green) and carotenoids (yellow-orange) absorb energy.
7. Light is absorbed by the electrons of the pigments (Figure 9.5); as a result, the electrons go from
ground state to an excited state.
8. When excited, electrons:
a. Return to their ground state by releasing their absorbed energy in the form of heat or fluorescence.
b. Are transferred to neighboring electron-accepting molecules called primary acceptor.
c. Pass their energy to the electrons of the neighboring molecules, making them the excited electrons.
C. Chlorophylls are the main light receptor pigments in photosynthesis.
1. Chlorophylls are the major photosynthetic pigments in plants, algae, and cyanobacteria.
2. Carotenoids act as accessory pigments whose electrons absorb light energy and pass it on to the
chlorophylls.
3. Chlorophyll has a magnesium atom at the center of the ring structure.
a. Chlorophyll a is found in plants, algae, and cyanobacteria.
b. Chlorophyll b is found only in plants and algae.
c. The absorption spectrum is the wavelength of light used by a pigment (Figure 9.7).

84 Chapter Nine
 d. The energy gathered by chlorophylls and carotenoids is relayed to chlorophyll a, which is the
main molecule directly involved in transforming light energy into chemical energy.
D. Photosynthetic pigments are organized into photosystems in chloroplasts (Figure 9.8).
1. Photosystems are embedded into the thylakoids.
2. Photosystems I and II collect different parts of the light reactions.
a. Each photosystem has an antenna, or light-harvesting complex.
b. It also has a reaction center where chemical energy is produced.
c. Photosystem I is called P700 due to the wavelength it absorbs.
d. Photosystem II is called P680 for the same reason.
E. Electron flow from water to NADP+ leads to the synthesis of NADPH and ATP.
1. Electrons are produced by the splitting of water: two per molecule of water.
2. Electrons are sent through a chain of electron carriers in photosystem II.
3. Then they are passed to photosystem I.
4. Energy from this transfer is used to create a proton gradient.
5. The proton gradient supplies energy to make ATP (Figure 9.10).
6. Then they pass through photosystem I and make NADPH.
7. NADPH supplies energy for CO2 fixation (Figure 9.10).
8. This is called linear electron flow.

Photosynthesis 85
 F. Electrons can also drive ATP synthesis by flowing cyclically around photosystem I (Figure 9.11).
1. Electrons flow through the cytochrome complex to P700 of photosystem I, excited by light energy,
then back to the cytochrome complex.
2. Electrons are not used for NADPH production.
3. Each cycle pumps one more proton pair across the membrane.
4. It is hypothesized that this pathway may be a physiological response to stress conditions, as ATP is
made but not NADPH. This limits reducing power and the consequent formation of reactive oxygen
species.
G. Experiments with chloroplasts confirm synthesis of ATP by chemiosmosis.
1. Jagendorf and Uribe placed chloroplasts in darkness first, then in acidic medium, and then into a basic
medium (Figure 9.10).
2. The change in proton gradient resulted in synthesis of ATP.

9.3 The Light-Independent Reactions of Photosynthesis

A. NADPH reduces CO2 into carbohydrates in the Calvin cycle.
B. The Calvin cycle uses NADPH, ATP, and CO2 to generate carbohydrates (Figure 9.12).
1. The dark reaction is separated into three phases.
a. Carbon from CO2 is added to ribulose 1,5-bisphosphate (RuBP) and produces two three-carbon 3-
phosphoglycerate (3PGA) molecules.
b. 3PGA is converted into G3P, another three-carbon molecule, which combines to form glucose.
c. Regeneration occurs using some G3P to produce RuBP.
2. Three turns of the Calvin cycle are needed to make one net G3P molecule (Figure 9.12).
a. From each turn of the Calvin cycle, two molecules of G3P are produced, which recycle in the
Calvin cycle.
b. Three turns of the Calvin cycle generate six molecules of G3P—one G3P goes toward the making
of glucose.
C. G3P is the starting point for synthesis of many other organic molecules.
1. Six turns of the Calvin cycle generate two G3P that will go toward making a molecule of glucose.
2. Once glucose is made, plants can form other molecules like sucrose, starch, and cellulose.
D. Rubisco is the key enzyme of the world’s food economy.
1. Rubisco is the first step of the Calvin cycle.
2. Rubisco converts 100 billion tons of CO2 into carbohydrates annually.

Focus on Basic Research: Two-Dimensional Paper Chromatography and the Calvin Cycle
A. Melvin Calvin and Andrew Benson of the University of California at Berkeley used radioactive carbon
dioxide to study dark reactions in green algae, Chlorella.
B. After exposing the cells to radioactive carbon dioxide for various amounts of time, they:
1. Stopped photosynthesis by placing the cells in alcohol.
2. Extracted the chemicals.
3. Put chemical samples on chromatography paper and ran them in two dimensions.
4. Put together chemicals in sequence to construct the Calvin cycle.

Insights from the Molecular Revolution: Small But Pushy: What Is the Function of the Small Subunit of Rubisco?
A. Rubisco has 16 subunits, eight large and eight small subunits.
B. The active site on rubisco is on the large polypeptide subunit.
C. If the small subunit is removed, it loses 99% of its function, but all the active sites seem to be on the large
subunits.
D. Betsy Reed and Robert Tabita of the Ohio State University—What is the function of small subunits?
E. Rubisco’s eight copies of each subunit all do the same thing—catalyze the same reaction.
F. The small subunits have a significant effect on rubisco’s rate of catalysis.

9.4 Photorespiration and Alternate Processes of Carbon Fixation

A. Photorespiration in plants is a mechanism similar to aerobic respiration that uses oxygen as the first step in
a pathway to generate CO2.
B. Photorespiration produces CO2 that is used by the Calvin cycle.
1. Rubisco has an active site to which either of the two gases (CO2 or O2) can bind.
2. When CO2 binds, rubisco is a carboxylase; when O2 binds, it is an oxygenase.
3. O2 is a competitive inhibitor, preventing CO2 from binding to the enzyme.

86 Chapter Nine
 4.During a hot day, O2 binds to rubisco, inhibits the enzyme, and reduces photosynthesis.
5.The upper surface of the leaves is covered by a cuticle that prevents water loss but also prevents rapid
diffusion of gases.
6. Most gas exchange occurs through stomata located on the under surface of the leaves.
7. Plants that are adapted or not adapted to hot, dry climates are faced with a constant dilemma: They
need to keep their stomata open for CO2 to diffuse, but closed to prevent water loss.
8. When stomata are closed, CO2 in the leaf is consumed and oxygen accumulates.
C. Alternate processes of carbon fixation minimize photorespiration.
1. Many plants that live in hot, dry environments have evolved alternate processes of carbon fixation that
minimize photorespiration, and therefore its negative effect on photosynthesis.
2. C4 plants and the C4 pathway.
a. C4 plants include corn, sugarcane, crabgrass, and many other members of almost 19 families of
flowering plants.
b. C4 plants have a characteristic leaf anatomy: photosynthetic mesophyll cells are tightly associated
with chloroplast-rich bundle sheath cells (bundle sheath cells are not photosynthetic in C3 plants)
(Figures 9.14a and 9.14b).
c. In C4 pathway, CO2 initially is fixed in the form of oxaloacetate by combining CO2 with PEP.
d. PEP is converted to malate and then pyruvate.
e. Pyruvate is oxidized to release CO2, which then is used by the Calvin cycle (Figure 9.15a).
f. Photorespiration is extremely low or negligible in C4 plants.
g. Why are all plants not C4? Because C4 plants have an additional requirement of ATP for each turn
of the C4 cycle.
3. CAM plants and the CAM pathway.
a. Just like C4 plants, CAM plants fix CO2 as oxaloacetate.
b. In CAM plants, carbon fixing and Calvin cycle both take place in mesophyll cells and at different
times of the day and night (Figure 9.15b).
c. CAM pathway takes place in cacti and orchids.

9.5 Photosynthesis and Cellular Respiration Compared

A. Photosynthesis and cellular respiration are similar processes, with reactions that essentially are the reverse
of each other (Figure 9.16).
1. Cellular respiration takes place in plants and in animals.
2. Photosynthesis occurs in plants, only in cells that contain chloroplasts.
3. Cellular respiration takes place in all the cells of plants.
4. The reactants of photosynthesis are products of cellular respiration and visa-versa.
5. Both photosynthesis and cellular respiration involve the electron transfer system—
photophosphorylation in photosynthesis and oxidative phosphorylation in cellular respiration—
followed by chemiosmotic synthesis of ATP.
6. The products of photosynthesis are the reactants of cellular respiration.
B. Evolution of photosynthesis and cellular respiration.
1. Earth’s early atmosphere had very little oxygen.
2. With the evolution of photosynthesis 2.7 billion years ago, oxygen levels rose.
3. At some point, prokaryotes developed aerobic respiration.
4. Through the process of endosymbiosis, both photosynthetic and aerobic prokaryotes were incorporated
into eukaryotic cells.

Unanswered Questions
A. Why is photosynthesis so inefficient, and what can we do about it?
1. Photosynthesis involves the highest energy processes of life.
2. The intermediates of photosynthesis are very reactive, to the point that much of the absorbed energy
must be dumped by plants.
3. A typical agricultural crop may store only about 0.1% to 0.5% of the absorbed energy into its biomass.
4. Some plants can store up to 10 times more energy.
B. How is the efficiency of photosynthesis regulated?
1. David Kramer of Washington State University is researching the energetics and regulation of
photosynthesis.
2. His laboratory has determined that production of heat helps to balance the production of damaging
components during photosynthesis.
3. ATP synthase acts as a major regulator of photosynthesis.

Photosynthesis 87
Suggestions for Presenting the Material
 One of the common student misconceptions that occurs regarding the cellular processes that generate ATP is that
plants do not undergo cellular respiration. Make sure that somewhere in the discussion of this chapter students are
reminded that Chapter 8 has relevance to photosynthetic organisms and that they also perform cellular respiration. Many
students also do not understand that for the Calvin cycle there are “three turns” of the wheel.
 Another common misconception is what the term carbon fixation really means. Do your best to help students realize
that when carbon is fixed, that means it is turned into new tissues in the organism. This is why many scientists are
concerned with plant growth for fixing carbon from the atmosphere.
 Many students have problems with considering the same enzyme rubisco as an oxygenase and a carboxylase.
Explaining to students how different reactions can be sequestered based on their location in the cell should help with this
difficulty.
 Review oxidation-reduction reactions again. After Chapter 8, these reactions should be very fresh in the students’
memory. Repetition of concepts like this can be helpful for many students in learning the reactions presented in
Chapter 9.
 A discussion of the reason colored objects appear the color they do may facilitate the students’ understanding of why,
for example, green light is ineffective for photosynthesis (not absorbed).
 Photosynthesis is much more complicated than the usual simple equation that a student may have seen in a previous
course. Perhaps the best approach to presenting this topic is to follow a stepwise outline and rely heavily on
transparencies and slides of the figures in the chapter. Giving handouts on which students have to fill in the products
from each step may help to reinforce learning the pathways.
 Refer to the old terminology of the light-dependent and light-independent reactions and why these are inaccurate.
 Explain to students the difference in the amount of ATP that can be generated from cyclic versus noncyclic pathways
in photosynthesis.
 Emphasize that plants do carry on aerobic respiration. Many students have the mistaken idea that plants only
photosynthesize.
 Stress to students the importance of knowing the processes rather than memorizing all the various reactions. Tell
students it is important to know what starts each step of the cycle and what the outcome of each cycle will be in order to
understand photosynthesis. For example: G3P is emphasized in the text as the starting point for the synthesis of many
other organic molecules.
 Talk about plants that are adapted to different types of habitats. Explain why there are C 3, C4, and CAM designations
for these plants. This will help students understand that there is not a single photosynthetic pathway, but even though
they all begin the same, there are differences that maximize efficiency in different habitats.
 Emphasize NADPH and ATP as carrier molecules. This reinforces Chapter 8 and shows the similarity of energy-
producing pathways across phyla.

Visual Learning Tips

 It may be useful to begin the lecture by showing the animation for the overview of photosynthesis. Giving students the
big picture and then breaking it down into parts helps many students categorize the different reactions of
photosynthesis.
 A slide of the global carbon cycle may be useful to increase student awareness of the importance of photosynthesis in
global biogeochemical cycles.
 Use Figures 9.4 and 9.7 to show different wavelengths of light and how different pigments absorb different
wavelengths.
 Use Figure 9.10 to show the orientation of ATP synthase facing out to the stroma, and the proton gradient being
formed in the thylakoid lumen. Also explain that, although the figure shows a linear and one-to-one arrangement of
Photosystem II–cytochrome complex–Photosystem I– NADP+ reductase–ATP synthase, there are fewer ATP synthase
along the membrane than this figure suggests.
 Make use of Figure 9.14 to illustrate the different anatomies of C4 and CAM plants. Again, stress the dual character of
rubisco and how plants have evolved to optimize photosynthetic reactions.

88 Chapter Nine
 Follow up your discussion of C3, C4, and CAM plants by showing the animation comparing C4 and CAM plants. For
visual learners, this can be a powerful tool.
 Have students “follow the carbon” through the Calvin cycle. This will help students understand the number of turns
necessary to produce one net G3P molecule. In addition, it will help them learn each step of the cycle if that is something
that will be emphasized in the classroom.

Classroom Discussion

Why It Matters
This section in the chapter highlights the discovery of the spectrum and organisms that perform photosynthesis. You can
use the BBC videos from the environmental science DVDs to give a global theme to the chapter as well. These can add a
poignant touch to the material covered and may result in a discussion about the importance of carbon fixation, rising
carbon levels in the atmosphere, and conservation. All of these topics can be touched on again during the conservation
chapters.

Focus on Research
Ask students to recall what they learned from the chemistry review in earlier chapters. Have them discuss why a method
like 2D paper chromatography was a useful method for Calvin and Benson.

Insights from the Molecular Revolution

Discuss the quantity of rubisco in animal cells and the rates of reaction. Ask students to come up with potential
explanations for why the small subunit of rubisco is so critical.

Unanswered Questions
Ask the students to discuss why plants have to regulate their rate of photosynthesis, and why they have to dissipate most
of their energy.

Additional Discussion Ideas

 Ask students why there is a dip in the absorption spectrum for green wavelengths (you can do this while showing
Figure 9.7).
 Discuss the importance of the antenna complex. Ask them to consider why each individual pigment molecule in the
antenna complex is unable to excite the photosystem on its own.
 Have students compare ATP synthase activity in mitochondria versus chloroplasts.
 Some students learn concepts better if presented with scenarios in which a pathway has “gone wrong.” Generate
discussion from students by asking what would happen if the stomata in CAM plants were unable to close completely
during the day when temperatures rise.
 The website http://photoscience.la.asu.edu/photosyn/study.html has great ideas for discussing the importance of
photosynthesis with students. The article “Why Study Photosynthesis” provides a good argument for calling
photosynthesis the most important biological process on Earth. The website also contains explanations for using reaction
centers of chloroplasts as microscopic models for exploring technologies that use light as an energy source.
 Usually we think of photosynthesis as producing carbohydrate and oxygen, but what important role does
photosynthesis perform as a “consumer”?

Classroom and Laboratory Enrichment

 Ask students to generate a concept map. You can give students a hint about what to include by referring students to
Figure 9.10.
 Have students make a table comparing and contrasting cyclic versus noncyclic electron flow.
 You can demonstrate the production of oxygen by plants with the following:
a. Place Elodea (an aquarium plant) in a bowl and expose it to bright light.
b. Invert a test tube over the plant and collect the bubbles.
c. Remove the tube and immediately thrust a glowing wood splint into the tube.

Photosynthesis 89
 d. Result: The splint burns brightly in the high-oxygen air.
 Separate the pigments in green leaves by using paper chromatography. (Consult a botany or biology laboratory manual
for the correct procedure.)
 Show an electron micrograph of a chloroplast and indicate where light-dependent reactions and light-independent
reactions occur.
 Cut out separated leaf pigments from a paper chromatogram and elute each pigment from the paper with a small
amount of alcohol. Using a spectrometer, determine the absorption spectrum for each pigment and graph the results.
(Consult a biology or botany laboratory manual for the procedure.)

Term Paper Topics, Library Activities, and Special Projects

 Students can go to the following website and look at a variety of animations and movies about ATP synthase:
http://www.csun.edu/~hcchm001/wwwatp2.htm.
 One of the hottest and driest summers in North America in recent years (1988) affected the production of food and
grain crops. Prepare an analysis of the effect(s) of that summer’s drought (or future ones) on future food availability and
its cost to the consumer.
 Consult a biochemistry or advanced plant physiology text to learn of a laboratory technique that would clearly indicate
whether the oxygen produced by plants is derived from water or from carbon dioxide.
 Prepare a detailed diagram of the Calvin-Benson cycle, showing the introduction of radioactively-labeled carbon
dioxide and its subsequent journey through several “turns” of the cycle.
 Of the total amount of sunlight energy impinging on a green plant, what percentage of the energy is actually converted
into glucose?
 Compare the mechanisms of C3 and C4 photosynthesis, and give examples of plants that fit into each category.
 Collect different plant samples from different habitats. Using paper chromatography, can you determine if the
pigments present in each plant are similar or different? Do you see differences between pigments found in different
habitats? Why or why not?
 Discuss why radiation with greater or lesser wavelengths than visible light is not generally used in biological
processes.

Suggested Readings

InfoTrac® College Edition Readings

 Gest, Howard. “Evolution of Knowledge Encapsulated in Scientific Definitions.” Perspectives in Biology and
Medicine. Autumn, 2001.
 Knight, Stefan, et al. “Reexamination of the Three-Dimensional Structure of the Small Subunit of RuBisCo from
Higher Plants.” Science. May 12, 1989.
 Wood, Marcia. “Robust Plants’ Secret? Rubisco Activase!” Agricultural Research. November, 2002.
 Pakrasi, Himadri B. “Genetic Analysis of the Form and Function of Photosystem I and Photosystem II.” Annual
Review of Genetics. 1995.
 Grossman, Arthur R., et al. “Light-Harvesting Complexes in Oxygenic Photosynthesis: Diversity, Control, and
Evolution.” Annual Review of Genetics. 1995.
 “Evolution of Photosynthesis.” Applied Genetics News. July, 2002.
 Vermaas, Wim. “Revealing the Secrets of Old Sol’s Sugar Factories.” World and I. March, 1998.
 Mann, Charles C. “Genetic Engineers Aim to Soup Up Crop Photosynthesis.” Science. January 15, 1999.
 Ben-Ari, Elia T. “CAM Photosynthesis: Not Just for Desert Plants.” BioScience. December, 1998.

Other Readings
The following is a website that students may enjoy visiting to learn more about the topics covered in Chapter 9.
http://www.cwnp.org/adaptations.html

90 Chapter Nine
Possible Responses to Think Outside the Book
The following website has a nice overview of research being done on harvesting energy from artificial photosynthesis:
http://www.scientificamerican.com/blog/post.cfm?id=shift-happens-will-artificial-photo-2010-03-03. You could direct
students to this site first so they can better refine their search. Their synopses of the research will likely include a
discussion on the splitting of water, how the sun’s energy can be used for splitting water, and some mechanisms for
harnessing the energy released when the chemical bonds of water are broken.

Possible Responses to Discuss the Concepts

1. Red, white, and blue petunias each absorb and reflect different wavelengths of light. The blue petunias absorb all
colors of the visible spectrum except blue, which they reflect, making them blue in color. The red petunias absorb all
colors except red, which they reflect. White petunias reflect all wavelengths, which combine to appear white.
2. Van Helmont’s experiment measured the trees and the soil; he may have even measured the weight of the water that
was added to the pots over the years. However, he didn’t know that the water that is given evaporates from the
leaves (transpiration), and that the carbon that is added to the biomass of the tree is actually taken from carbon
dioxide in the atmosphere.
3. From the evidence given, it seems that the creation of oxygen-free radicals by chlorophyll molecules leads to the
destruction of these molecules. Without antioxidant mechanisms, the plants will appear bleached and may
eventually die as they lose chlorophyll and, therefore, their ability to photosynthesize.
4. In order for a plant to produce glucose in the dark it needs carbon dioxide, NADPH, ATP, and, most importantly, the
enzyme rubisco. In photosynthesis, ATP and NADPH are generated in the light-requiring reactions. Glucose can be
produced in the dark, as long as light reactions precede dark reactions.

Possible Response to Design an Experiment

To determine if the purple beings (PB) harvested energy by photosynthesis, the researchers should first determine if PB
ingests organic compounds. If they do, these compounds are likely these organisms’ source of energy as well as carbon.
If they do ingest carbon compounds, they may still be photoheterotrophs—getting their energy from sunlight, but their
carbon from preformed carbon compounds. The easiest way to determine if the PB are photosynthetic is to remove them
from natural sunlight. If they are photosynthetic, they will eventually get “hungry” as their stores of ATP and NADPH
are depleted due to the lack of light-energy influx.

Possible Response to Apply Evolutionary Thinking

If global warming continues, C4 plants that are more dry-adapted will be favored over C3 plants. The distribution of
deciduous trees will increase, and the taiga (coniferous forest) will decrease. The cold and dry tundra regions will also
shrink.

Photosynthesis 91