Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Color It Green
ts easy to show that plants do not use green light for photosynthesis. Simply put a sprig of the plant elodea in a glass jar, ll it with water (elodea lives in water), and shine a green light on it. NOTHING HAPPENS. But switch to a bright white light, and watch the bubbling. Bubbling is caused by oxygen gas escaping from the water. Thats your evidence that photosynthesis is occurring. Plants always give off oxygen when they photosynthesize, for which we humans are mightily thankful. White light, the visible light that shines down on us every day, contains different colors of light, from violet to blue, green, yellow, orange, and nally red. Plants use all the colors except greenand thats why we see them as green! Red algae are protists that live in the ocean, and like all the other types of algae, they photosynthesize. Despite their name, some forms of red algae are
APPLI C AT I O N S
HOW LIFE CHANGES
104 1 10 04
dark-colored, almost black, which means that they are able to use all the different colors in white light for photosynthesis. Does this mean that if plants werent so wasteful and used green light for photosynthesis, in addition to all the other colors, they would appear black to us? Yes, it does. Look out the window and imagine that all the plants you see are black instead of green. Arent we glad that photosynthesis on land is inefcient and wasteful of green light, making our world a sea of green! How did it happen that plants do not use green light for photosynthesis? During the evolution of organisms, photosynthesizing bacteria oating in the oceans above the sediments possessed a pigment that could absorb and use green light. Natural selection, therefore, favored the evolution of a pigment that absorbed only blue and red light. The green pigment chlorophyll, which absorbs the blue and red ranges of light, evolved and became the
photosynthetic pigment of plants. The inefciency of chlorophyll doesnt matter on land, where light is readily available, so no plants on land ever evolved a more efcient pigment than chlorophyll. Plants are green and our world is beautiful because their primary photosynthetic pigment doesnt absorb green light! Green leaves and variously colored eukaryotic algae carry on photosynthesis in chloroplasts, as discussed in this chapter. Photosynthesis, which produces food in the form of a carbohydrate and also oxygen for the biosphere, consists of two connected types of metabolic pathways: the light reactions and the Calvin cycle reactions. We will see how the absorption of solar energy during the light reactions drives the Calvin cycle reactions, that produce a carbohydrate, the end product of photosynthesis. This chapter also shows that the process of photosynthesis is adapted to different environmental conditions.
105
List the types of organisms that carry on photosynthesis. (6.1) Identify the main parts of a chloroplast, a eukaryotic organelle. (6.2) Write a redox equation for photosynthesis that shows both the reactants and the products. (6.3) Divide photosynthesis into two sets of reactionsone set that captures solar energy and the other that reduces carbon dioxide. (6.3)
Photosynthesizers not only produce food for the biosphere, but they are also the source of the fossil fuel our society burns to maintain its standard of living. The overall equation for photosynthesis shows the starting reactants and the end products. But we will see that, in actuality, photosynthesis requires two metabolic pathways: the light reactions that occur in thylakoid membranes and the Calvin cycle reactions that occur in the stroma of a chloroplast.
Euglena, a protist
kelp, a protist
moss, a plant
mesophyll
3 CO2
2 leaf vein
stroma granum
stroma
4 Chloroplast
37,000
107
6.3 Photosynthesis involves two sets of reactions: the light reactions and the Calvin cycle reactions
During photosynthesis, hydrogen atoms are transferred from water to carbon dioxide with the release of O2 and the formation of glucose:
Reduction solar energy
CO2 +H2O
(CH2O)+O2
The energy of these electrons empowered by the sun is captured and later used for ATP production. Energized electrons are also taken up by a coenzyme called NADP+ (nicotinamide adenine dinucleotide phosphate). After NADP+ accepts electrons, it combines with an H+ derived from water. The reaction that reduces NADP+ is: NADP++2 e+H+ NADPH The lower set of red arrows in Figure 6.3 show that the NADPH and ATP produced by the light reactions are sent to the Calvin cycle reactions.
Oxidation
In chemistry, oxidation is the loss of electrons, and reduction is the gain of electrons. In cells, oxidation is instead the loss of hydrogen atoms, and reduction is the gain of hydrogen atoms. This difference is easily explained because a hydrogen atom contains one electron and one hydrogen ion (e+H+); when a molecule loses a hydrogen atom, it has lost an electron, and when a molecule gains a hydrogen atom, it has gained an electron. Therefore, photosynthesis is indeed an oxidation-reduction, usually shortened to a redox reaction. Even though we often use the overall reaction for photosynthesis, researchers have known for some time that photosynthesis involves two sets of reactions. These two sets of reactions can be associated with the two parts of a chloroplastnamely, the stacks of thylakoids (grana) and the stroma, as shown in Figure6.3.
Light Reactions The light reactions (sometimes called the lightdependent reactions) are so named because they only occur when solar energy is available (during the daylight hours). At that time, the chlorophyll molecules and other pigment molecules located within the thylakoid membranes absorb solar energy and use it to energize electrons taken from water, which splits, releasing oxygen: H2O e +2H + O2
+
Calvin Cycle Reactions The Calvin cycle reactions (sometimes called the light-independent reactions because they can occur both day and night) are named for Marvin Calvin who discovered them and their cyclical arrangement. The enzymatic reactions of the Calvin cycle occur in the stroma of a chloroplast. During the Calvin cycle, CO2 is taken up and then reduced to a carbohydrate that can be used to form glucose. Reduction of CO2 to a carbohydrate (CH2O) requires hydrogen atoms supplied by NADPH and energy supplied by ATP, both molecules provided by the light reactions. The upper set of red arrows in Figure 6.3 show that after the Calvin cycle reactions are complete, ADP + P and NADP+ are sent back to the light reactions.
6.3 Check Your Progress What two molecules form a bridge
linking the light reactions with the Calvin cycle reactions, and what molecules function similarly in the opposite direction?
CO2
stroma
108
PART 1
Explain why leaves are green, with reference to the electromagnetic spectrum. (6.4) Explain how solar energy is converted to that of ATP during photosynthesis. (6.56.6) Trace the path of excited electrons, from the absorption of solar energy to the production of ATP and NADPH. (6.7) Explain how the thylakoid is organized to produce ATP and NADPH. (6.8)
The light reactions begin when pigments in the thylakoid membrane absorb certain wavelengths of light. The thylakoid membrane is highly organized for absorbing solar energy and producing ATP and NADPH, needed by the Calvin cycle reactions to reduce carbon dioxide to a carbohydrate in the stroma.
Increasing wavelength chlorophyll a chlorophyll b carotenoids Increasing energy Gamma rays X rays UV MicroInfrared waves Radio waves Relative Absorption 380
visible light
380
500
600
750
500
600
750
Wavelengths (nm)
Wavelengths (nm)
109
FIGURE 6.5
A general model of a photosystem.
energy level electron acceptor
e:
solar energy
e: reaction center
antenna molecule
pigment complex
6.5 Check Your Progress What is the function of antenna molecules in the pigment complex of a photosystem?
low-energy electrons
e:
6.6 Check Your Progress What kind of energy is the ow of hydrogen ions down their concentration gradient through an ATP synthase?
110
PART 1
CO2
sun
Light reactions
sun
NADPH ATP
electron acceptor
electron acceptor
energy level
e: e
:
ele
ctro
e: NADP+
n tr ans
por
ATP
7
t ch
ain
e:
H+
e: reaction center
1
10
NADPH
reaction center
pigment complex
4
pigment complex PS I e
:
PS II CO2
5
H2O
6
CH2O
2 H+
1 2
O22
FIGURE 6.7 During the light reactions, electrons follow a noncyclic pathway in the thylakoid membrane; electrons move from water to NADP+.
CHAPTER 6 Pathways of Photosynthesis
111
Application
F O R M YO U R O P I N I O N
1. Evolution proceeds by adding on rather than starting over. How is the evolution of oxygen-releasing photosynthesis an example of this concept? 2. Often during evolution, attributes are lost. Today, what ability do free-living cyanobacteria have that chloroplasts do not have?
sun
energy level
electron acceptor e:
ele ctr o
ATP
nsp tra
or t chain
e: reaction center
112
PART 1
Payoff
6
Get Ready
1
PS II consists of a pigment complex that absorbs solar energy and passes electrons on to an electron-acceptor molecule. 2 PS II receives replacement electrons from water, which splits, releasing H+ and oxygen (O2). The electron transport chain, consisting of a series of electron carriers such as cytochrome complexes, passes electrons from PS II to PS I. (Notice, therefore, that PS I receives replacement electrons from the electron transport chain.) 4 Members of the electron transport chain also pump H+ from the stroma into the thylakoid space. Eventually, an electron gradient is present: The thylakoid space contains much more H+ than the stroma. PS I consists of a pigment complex that absorbs solar energy and sends excited electrons on to an electron-acceptor mol-
NADP+ reductase, an enzyme, receives electrons and reduces NADP+. NADP+ combines with H+ and becomes 7 NADPH. H+ ows down its concentration gradient through a channel in an ATP synthase complex. This complex contains an enzyme that then enzymatically binds ADP to P , producing 9 ATP.
This method of producing ATP is called chemiosmosis because ATP production is tied to an H+ gradient across a membrane. We have now concluded our study of the light reactions. In the next part of the chapter, we will study the Calvin cycle reactions. 6.8 Check Your Progress What is the end result of the get ready
phase, and what is the end result of the payo phase of the light reactions?
thylakoid
thylakoid space
PS II H+
1
stroma H+
7
PS I Pq e: e:
4 2
NADP reductase e: e:
NADPH NADP+
H 2O solar energy CO2
e: H+ H+ O2
H2O
2 H+ +
1 2
NADPH ATP
H+
H+
ATP synthase
9
thylakoid membrane
stroma O2 CH2O
Thylakoid space
113
From the overall equation for photosynthesis, associate carbon dioxide and carbohydrate (CH2O) with the Calvin cycle reactions. The Calvin cycle reactions are light-independent because they can occur in the light or the dark, but even so they could not occur without the ATP and NADPH from the light reactions because they are needed to reduce CO2 to a carbohydrate.
6.9 ATP and NADPH from the light-dependent reactions are needed to produce a carbohydrate
The Calvin cycle is a series of reactions that produces carbohydrate before returning to the starting point once more (Fig. 6.9). Melvin Calvin and his colleagues used the radioactive isotope 14 C as a tracer to discover its many reactions. The reactions can be divided into these phases: CO2 xation, CO2 reduction, and nally regeneration of RuBP, the molecule that combines with and xes CO2 from the atmosphere. The steps in the Calvin cycle are multiplied by three for reasons that will be explained.
H2O solar energy CO2
CO2 Fixation
1 During the rst phase of the Calvin cycle, CO2 from the atmosphere combines with RuBP, a 5-carbon molecule, and a C6 (6-carbon) molecule results. The enzyme that speeds this reaction, called RuBP carboxylase, is a protein that makes up 2050% of the protein content in chloroplasts. The reason for its abundance may be that it is unusually slow (it processes only a few molecules of substrate per second compared to thousands per second for a typical enzyme), and so there has to be a lot of it to keep the Calvin cycle going.
ADP+ P
NADP+
Light reactions
NADPH ATP
3 CO2
stroma O2 CH2O
intermediate
3 C6
2
3PG C3
6 ATP
3
3 ADP + 3 P
5
Calvin cycle
CO2 reduction
6 ADP + 6 P
These ATP and NADPH molecules were produced by the light reactions.
RuBP regeneration
6 BPG C3
3 ATP
4
6 NADPH
5 G3P C3 6 G3P C3 net gain of one G3P
6 NADP+
x2 Glucose
114
PART 1
The C6 molecule immediately splits into two C3 molecules. (Remember that all molecules are multiplied by three in Figure 6.9.) This C3 molecule is called 3PG.
CO2 Reduction
3 and 4 Each of the 3PG molecules undergoes reduction to G3P in two steps:
allow one G3P to exit. Why? Because, for every three turns of the Calvin cycle, ve molecules of G3P are used to re-form three molecules of RuBP, and the cycle continues. Notice that 5 3 (carbons in G3P) = 3 5 (carbons in RuBP):
5 G3P
ATP
ADP+ P
3 RuBP 3 ADP+3 P
3 ATP
3PG BPG G3P
As this ATP produced by the light reactions breaks down, ADP+ P results.
NADPH
NADP+
During this phase, CO2 is reduced to a carbohydrate because RCO2 (3PG) has become RCH2O (G3P). Energy and electrons are needed for this reduction reaction, and these are supplied by ATP and NADPH, which become ADP+ P , and NADP+, respectively.
The ADP+ P and NADP+ that result from the Calvin cycle return to the light reactions so that they can continue to produce ATP and NADPH for the light-independent reactions. 6.9 Check Your Progress Justify the use of the expression lightindependent reactions for the Calvin cycle even though it is still very much dependent on solar energy. Explain.
RuBP Regeneration The reactions in Figure 6.9 are multiplied by three because it takes three turns of the Calvin cycle to
G3P
7 1
glucose phosphate
+ fructose phosphate
sucrose
starch
cellulose
FIGURE 6.10 Trees and all photosynthesizers have the metabolic pathways that are able to convert G3P to all the molecules noted.
CHAPTER 6 Pathways of Photosynthesis
115
Thus far, we have been observing C3 photosynthesis, named for the number of carbons in the rst observable molecule following the uptake of CO2. In contast, C4 photosynthesis is more advantageous when the weather is warm and CO2 is in short supply inside leaves, due to closure of stomata. CAM, a third type of photosynthesis, was rst observed in desert plants, which need to conserve water.
3PG (C3)
combining with O2. As we shall see in Section 6.12, C4 plants have such an advantage when the weather is warm. 6.11 Check Your Progress Explain the term C3 photosynthesis.
C3 Plant
C4 Plant
mesophyll cells
vein stoma
vein stoma
116
PART 1
In C4 plants, the Calvin cycle reactions occur in the bundle sheath cells and not in the mesophyll cells. Therefore, CO2 from the air is not xed by the Calvin cycle. Instead, CO2 is xed by a C3 molecule, and the C4 that results is modied and then pumped into the bundle sheath cells (Fig. 6.12B). Now the C4 molecule releases CO2 to the Calvin cycle. This represents partitioning (of pathways) in space. It takes energy to pump molecules, and you would expect the C4 pathway outlined in Figure 6.12B to be disadvantageous. Yet, in warm climates when stomata close and CO2 is in limited supply inside leaves, the net photosynthetic rate of C4 plants (e.g., sugarcane, corn, and Bermuda grass) is two to three times that of C3 plants (e.g., wheat, rice, and oats). Why do C4 plants enjoy such an advantage? The answer is that RuBP carboxylase located in bundle sheath cells always combines with CO2 and does not combine with O2. When the weather is moderate, C3 plants ordinarily have the advantage, but when the weather becomes warm C4 plants have their chance to take over, and we can expect them to predominate. In the early summer, C3 plants such as Kentucky bluegrass and creeping bent grass are predominant in lawns in the cooler parts of the United States, but by midsummer, crabgrass, a C4 plant, begins to take over.
Calvin cycle
G3P
Section 6.13 discusses still another form of photosynthesis, called CAM photosynthesis, which is prevalent in desert plants. 6.12 Check Your Progress Structure suits function. How do C4 plants prevent exposure of the RuBP carboxylase enzyme to oxygen?
day
G3P
Where the climate is hot and dry (deserts), stomata close up during the day and CAM plants become competitive also. The occurrence of climate change is accompanied by an increased level of CO2. How Science Progresses on page 118 discusses how tropical rain forests can help prevent any further rise in atmospheric CO2. 6.13 Check Your Progress How is CAM photosynthesis partitioned by the use of time?
CHAPTER 6
Pathways of Photosynthesis
117
Application
F O R M YO U R O P I N I O N
1. What are some other advantages to preserving tropical rain forests aside from helping to prevent any further rise inCO2? 2. What can the countries of the world that have no tropical rain forests do to help preserve them?
5.5 Mean Global Temperature Change (C) 4.5 3.5 2.5 1.5 0.5 atmospheric CO2
FIGURE 6B This graph shows how atomospheric CO2 is expected to aect global temperature. As shown the global temperature is now 0.05C higher world wide than in the 20th century. Burning forests adds to atmospheric CO2.
118
PART 1
CO2 +H2O
(CH2O)+O2
Oxidation
The light reactions only occur in thylakoids during the day when solar energy is available. The light reactions split water, releasing O2, and send NADPH and ATP to the light reactions. The Calvin cycle reactions are enzymatic reactions that occur in the stroma anytime, day or night. The Calvin cycle reactions use the NADPH and ATP from the light reactions to reduce CO2 to a carbohydrate (CH2O).
CO2
NADPH ATP
stroma
119
CO2 reduction: Each 3PG is reduced to a G3P molecule. This step requires ATP and NADPH:
ATP
3PG ADP+ P BPG NADPH G3P NADP+
c. convert light energy into usable forms of chemical energy. d. regenerate RuBP. 5. Label the following diagram of a chloroplast.
H2O solar energy ADP + P NADP+ CO2
RuBP regeneration: During three turns of the Calvin cycle, ve molecules of G3P are used to re-form three molecules of RuBP. This step requires ATP energy. 6.10 In photosynthesizers, a carbohydrate is the starting point for other molecules G3P (phosphoglyceraldehyde-3-phosphate) can be converted to all organic molecules needed by a plant. It takes two G3P molecules to make one glucose molecule.
d.
Light reactions NADPH ATP
a.
b. c.
CH2O
e.
6. THINKING CONCEPTUALLY In order for the biosphere to have animal life, some animals must eat plants. Explain.
TESTING YOURSELF
Photosynthesis Produces Food and Releases Oxygen
1. The raw materials for photosynthesis are a. oxygen and water. b. oxygen and carbon dioxide. c. carbon dioxide and water. d. carbohydrates and water. e. carbohydrates and carbon dioxide. 2. During photosynthesis, __________ is reduced to __________. a. CO2, oxygen c. water, oxygen b. oxygen, CO2 d. CO2, a carbohydrate 3. The light reactions a. take place in the stroma. c. Both a and b are correct. b. consist of the Calvin cycle. d. Neither a nor b is correct. 4. The function of the light reactions is to a. obtain CO2. b. make carbohydrate.
120
PART 1
d. use the products of the light reactions. e. All of these are correct. 14. The Calvin cycle requires ______ from the light reactions. a. carbon dioxide and water d. ATP and water b. ATP and NADPH e. NADH and water c. carbon dioxide and ATP 15. THINKING CONCEPTUALLY The overall equation for photosynthesis doesnt include ATP. Why is ATP needed?
18. The different types of photosynthesis are dependent upon the timing and location of a. CO2 xation. c. H2O xation. b. nitrogen xation. d. All of these are correct.
THINKING SCIENTIFICALLY
1. Elodea, a plant that lives in the water, is in a beaker of water. Bubbling occurs with white light, but not green. Why do environmental conditions have to be kept constant, and suggest a control for this experiment. 2. The process of photosynthesis supports the cell theory. How?
ONLINE RESOURCE
www.mhhe.com/maderconcepts2
Enhance your study with animations that bring concepts to life and practice tests to assess your understanding. Your instructor may also recommend the interactive eBook, individualized learning tools, and more.
CHAPTER 6
Pathways of Photosynthesis
121