CDN ED Biology Exploring the Diversity of Life 2nd Edition

Russell
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7 Photosynthesis
IF NOTHING ELSE, MY STUDENTS SHOULD LEARN…

1. Photosynthesis is arguably the most important biochemical process on Earth.

2. Photosynthesis is a set of reactions that capture light energy into organic molecules
that can be broken down for energy (by the plants themselves, as well as
heterotrophs).

LEARNING OBJECTIVES

Students should be able to:

 Define key terms: chlorophyll, mesophyll, stomata, primary acceptor, accessory

pigments, absorption spectrum, action spectrum, photosystem I (PSI), photosystem II
(PSII), antenna complex, reaction centre, P700, P680, water-splitting complex,
photophosphorylation, noncyclic photophosphorylation, cyclic photophosphorylation,
Ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco), carbon-concentrating
mechanism, photorespiration, phosphoenolpyruvate (PEP), bundle sheath cells.
[Knowledge]

 List and describe the two parts/stages of oxygenic photosynthesis in general terms.

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-1

 [Comprehension]

 Describe the major plant structures involved in photosynthesis in a green plant (i.e.,
organ, tissue, organelle). [Comprehension]

 Describe the structure of the chloroplast. [Knowledge]

 Explain how the membranes/compartments of chloroplast (structure) relate to the

processes of photosynthesis (function). [Comprehension, Analysis]

 List and briefly describe the pigments involved in photosynthesis. [Comprehension]

 Explain why there are different pigments in an antenna complex. [Comprehension]

 Describe the possible fates of an excited-state electron in a pigment molecule.

[Comprehension]

 Relate the absorption spectrum of photosynthetic pigments in a given plant with the
action spectrum of photosynthesis in that plant. [Comprehension, Application]

 Correctly interpret a figure showing absorption spectra of photosynthetic pigments

and/or action spectrum of photosynthesis in a given plant. [Application, Analysis]

 Describe, in general terms, what occurs in light-dependent reactions of

photosynthesis (including energy investments, energy yields, oxygen, water and
carbon inputs/outputs). [Comprehension]

 Describe the basic structure, components, and function of photosystems, and the
roles of PSI and PSII in the light-dependent reactions. [Comprehension]

 Describe how ATP is produced in the light-dependent reactions. [Comprehension]

 Explain the role of light for electron transfer in photosystems. [Comprehension]

 Compare and contrast linear electron transport and cyclic electron transport.
[Analysis]

 Describe (briefly) the following stages of the Calvin cycle: CO2 fixation; production
of G3P; regeneration of RuBP. [Comprehension]

 Describe Rubisco, discussing both the carboxylase and oxygenase activities of this
enzyme. [Comprehension]

 Describe, in general terms, what occurs in the light-independent reactions/Calvin

cycle (including energy investments, yields and carbon inputs/outputs).
[Comprehension]

7-2 CHAPTER 7 Copyright © 2013 by Nelson Education Ltd.

  Describe ribulose 1,5-bisphosphate (RuBP), 3-phosphoglycerate, and glyceraldehyde-
3-phosphate (G3P) in terms of carbons/phosphates and role in the Calvin cycle.
[Comprehension]

 Describe the importance of Rubiscoto life on Earth. [Comprehension]

 Correctly predict the outcome in cellular respiration or photosynthesis if a particular

key molecule (e.g., enzyme, organelle structure) is damaged/missing/inhibited.
[Application]

 Describe photorespiration, discussing the conditions that lead to this process

occurring. [Comprehension]

 Explain how photorespiration is different from cellular respiration. [Comprehension,

Analysis]

 Describe the challenges faced by, and strategies employed by aquatic

photosynthesizers in terms of obtaining enough CO2. [Comprehension]

 Describe how the C3, C4, and CAM pathways of carbon fixation differ, indicating the
advantages and disadvantages of each pathway. [Comprehension]

 Compare substrate level phosphorylation, oxidative phosphorylation (discussed in

Chapter 6), and photophosphorylation. [Comprehension, Analysis]

 Identify various cellular processes, including biochemistry, energetics, necessary

proteins, and membranes that are similar in cellular respiration and photosynthesis.
[Analysis, Synthesis]

WHY IS THIS CHAPTER IMPORTANT TO SCIENTISTS?

 Photosynthesis is an ancient, important set of biochemical reactions.

 Herbicide development is often directed at interfering with photosynthesis in target

weeds.

 Climate change has been linked to substantial increases in greenhouse gases (e.g.,
CO2) as a result of human activity. Photosynthesis is a natural carbon sink (i.e., stores
carbon compounds) that removes CO2 from the environment and can be used as a
carbon offset. Researchers are studying ways of countering the effects of climate
change with photosynthesizing plants.

 The nature of some of the components of photosynthesis may have applications in

other areas (e.g., photochemical switches, improved sunscreens, cancer therapies).

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-3

WHY SHOULD STUDENTS CARE?

 Nearly all life on Earth depends (directly or indirectly) on the products of

photosynthesis.

 Climate change has been linked to substantial increases in greenhouse gases (e.g.,
CO2) as a result of human activity. Understanding the role of photosynthesis in
capturing carbon can help students understand the importance of photosynthesizers
in the environment.

 Elements of photosynthesis are examples of biochemical reactions that appear in

other reactions; an understanding of photosynthesis will help students with other
concepts.

WHAT ARE COMMON STUDENT

MISCONCEPTIONS/STUMBLING BLOCKS?

1. Students may have difficulty realizing that “photosynthesis” refers to many individual
reactions, occurring in sets. It is recommended that instructors explicitly mention
that the term “photosynthesis” is generally used as shorthand to encompass the sum
of all the reactions occurring in different stages (and locations in the chloroplast).

2. We have found that many students are focused on simply memorizing steps/
enzymes for the reactions, without considering the “bigger picture” of what is going
on during these processes. Clear expectations from the instructor regarding what
students are expected to know may help alleviate some of the anxiety associated with
massive memorization (if, indeed, the instructor does not want students to memorize
all of this information).

3. The light-independent reactions (Calvin cycle) are sometimes referred to as the

Calvin-Benson Cycle, or the dark reactions. The latter is most problematic, as the
reactions are not restricted to the dark (while light reactions are, indeed, only
possible when light is available).

4. Stroma (the chloroplast “cytoplasm” containing enzymes of the Calvin cycle) and
stoma (singular; plural = stomata; pores in plant leaves allowing gas exchange) are
words that students often mix up. These terms should be explicitly defined, and used
often if you expect students to know them both.

5. If students have already learned about cellular respiration (Chapter 6), they may find
the stages of photosynthesis confusing. ATP production occurs in all the major
stages of cellular respiration (glycolysis, Krebs cycle, ETC/oxidative
phosphorylation), but only in the light reactions of photosynthesis. This makes sense
if one understands that the CO2 fixation reactions (which are necessary for
production of chemical compounds used for longer-term energy storage and other

7-4 CHAPTER 7 Copyright © 2013 by Nelson Education Ltd.

 metabolic pathways) are extremely energy-intensive. Energy is provided via photons,
but must be converted into molecules that the Calvin cycle reactions can use.

6. The idea that three CO2 molecules enter the Calvin cycle is often nonintuitive to
students, as is the idea that several steps are required just to regenerate the “starting
molecule” (RuBP). Following carbon molecules through the cycle can help students
better visualize this.

7. The dual carboxylase and oxygenase activity of Rubisco can be extremely

counterintuitive. Context regarding the origin, in anoxic conditions before
appearance of oxygen-generating cyanobacteria, and long evolutionary history of this
enzyme may help with understanding this concept.

8. Students may have preconceived notions that they will not be interested in plants
(and that photosynthesis is not relevant to them), which may mean that there are
challenges in engaging them with the material. (They also may not have much
background in terms of understanding the structure of plants, including leaf
anatomy.) Highlighting the importance of photosynthesis (in supporting nearly all
life forms) may help, along with some of the environmental and applied aspects of
photosynthesis research.

9. Students may be unaware that photosynthesis occurs in many other types of

organisms beyond plants (e.g., some bacteria, archaeans, algae). Instructors may want
to remind students of Chapter 3, where the activity of ancient cyanobacteria led to an
oxygenated atmosphere.

10. In introductory biology, we often present only cellular respiration and

photosynthesis as examples of metabolic pathways. This can lead students to have a
narrow view of what metabolism involves. Separate chapters for cellular respiration
and photosynthesis can also give the impression that these pathways are extremely
different topics, rather than examples of major metabolic pathways.

WHAT CAN I DO IN CLASS?

The “Why It Matters” introduction highlights the scale of the CO2 fixation that is
accomplished by photoautotrophs on Earth. This could be used to lead into discussion
about the importance of CO2 fixation, and issues that we are facing with regards to global
biodiversity and climate change issues. The nature and importance of phytoplankton can be
highlighted.

Ask students to consider the term “photosynthesis” in terms of “photo-” and “-synthesis,”
relating these parts of the word to the different stages of reactions that occur.

Revisit Chapter 1 (Light and Life), reviewing the nature of photosynthetic pigments, and
how the structure of these molecules is important in capturing energy from light. It may also

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-5

be valuable to revisit portions of Chapter 2 (The Cell: An Overview), particularly the
chloroplast.

Instructors may want to bring in actual plants to class to demonstrate the regions where
photosynthesis occurs, and adaptations that allow plants to obtain the water, oxygen, and
minerals they require. Using dissecting microscopes, aspects of leaf anatomy can be
examined (in a small class or lab; using a microscope camera for larger demonstrations).
Prepared slides may be useful to show the cells and tissues within leaves. Electron
micrographs can be shown when discussing chloroplast structure. [Stumbling Block 8]

If cellular respiration (Chapter 6) has already been covered, ask students to identify where
there are similarities and differences between photosynthesis and cellular respiration.
[Stumbling Block 10] You may want to provide students with a blank summary table to help
guide them with this task:

Photosynthesis Cellular Respiration

Overall reaction (simplified)

Reactants

Products

Cellular location(s)

Energy required? Released?

Stages/phases

Summary (one sentence

description)

Students can also be encouraged to draw figures representing mitochondria and

chloroplasts, and indicate where major phases of the reactions occur, including the electron
transport chain, and chemiosmosis (e.g., Where do we see a high concentration of H+?).
[Stumbling Block 10]

Instructors are advised to explore the events of the light reactions with students using
Figure 7.11 or an animation (e.g., from CengageNOW) to help students understand the steps
that occur in this process, similarities to some events in cellular respiration (reduction of an

7-6 CHAPTER 7 Copyright © 2013 by Nelson Education Ltd.

electron carrier, ETC, development of a proton gradient, chemiosmotic generation of ATP)
as well as the differences (light is needed to drive the transport of electrons in
photosynthesis, NADP+ vs. NAD+, etc.).

The animations of aspects of photosynthesis will be valuable to illustrate the important

steps that occur. The Calvin cycle animation may help students with Stumbling Block 6.

Rubisco can be explored in terms of the structure and function of the enzyme, and the fact
that the enzyme has both carboxylase and oxygenase activity. Ask students when in Earth’s
history Rubisco would have appeared, and what the conditions would have been like at that
time. [Stumbling Block 7] You may want to refer back to Chapter 3, and discuss the
appearance of oxygenic photosynthesizers and importance of this to the life that evolved
after significant amounts of O2 accumulated in the atmosphere.

An activity can be done in class, with students considering and comparing the benefits and
costs of C3, C4, and CAM photosynthesis under different conditions of temperature and
humidity. (In order to do this, students will need to know what the differences are between
these pathways.) If class size permits, teams can be assigned a type of photosynthesis to
analyze. See clicker question 5.

Example C4 and CAM plants can be brought into class to demonstrate the different types
of plants that carry out the variations of photosynthesis (e.g., C4—tropical grasses, maize,
sedges; CAM—succulent cacti, such as jade plants). Explicitly describing and highlighting
the differences in leaf anatomy between C3 and C4 plants in class (with a diagram or
microscope picture) can be valuable, especially if students are not very familiar with plant
cell/tissue types.

In a discussion or one-minute paper activity, ask students to consider challenges that are
faced by aquatic photosynthesizers. (This can lead into exploring the carbon-concentrating
mechanisms in algae.)

We think of sugar as the product of photosynthesis, but it should be highlighted that the
final storage product of photosynthesis is typically starch in most plants. (Sugars are
converted to starch.) Sugar cane and sugar beets are unusual in storing the sugar sucrose.
Plant-breeding efforts increased the sugar content in the beets from 2–6 percent to the
approximately 20 percent seen today. (This is an example of artificial selection.)

The traditional textbook chapter order places cellular respiration (Chapter 6) before
photosynthesis, and many instructors will follow this progression. It is possible to combine
concepts from these chapters (potentially with all or part of Chapter 4) to discuss
metabolism. In this way, the instructor and students can explore similarities (e.g., electron
transport chains, chemiosmosis, cycles that require regeneration of intermediate molecules,
etc.) as well as the differences between different types of metabolic pathways. Additional
metabolic pathways could also be introduced—in the past, we have shown the folate
pathway, as there are links to cancer (which relates to material in subsequent chapters).
[Stumbling Block 10]

Clicker questions:

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-7

 1. In green plants, which of the following occur(s) during the light-dependent
reactions?

A. Splitting of water into oxygen and hydrogen ions

B. reduction of NADP+
C. Transport of electrons along an electron transport chain
D. chemiosmotic production of ATP
E. all of the above *

This is a fairly straightforward question addressing the major activities of the light reactions.
[Stumbling Block 2]

2. The reactions of the Calvin cycle

A. can only occur in the light.

B. can only occur in the dark.
C. can only occur when there are adequate quantities of CO2, ATP and NADPH. *
D. both B and C

This question can identify if students have the misconception that the Calvin cycle only
occurs in the dark (as could be inferred from the term “dark reactions,” as this cycle is
occasionally called). [Stumbling Block 3]

3. Glyceraldehyde 3 phosphate (G3P)

A. is an intermediate in the Calvin cycle.

B. is a byproduct in the tryptophan biosynthetic pathway.
C. is a reactant in the thiamine biosynthetic pathway.
D. can be combined and rearranged to form other carbohydrates (e.g., glucose,
starch).
E. All of the above are true of G3P. *

This is a question intended to highlight that metabolic pathways do not operate in isolation.
We often see G3P indicated simply as the end-product of the Calvin cycle, but there are
other metabolic roles with which it is involved. [Stumbling Block 10]

4. Photosynthesis occurs in

A. plants only.
B. plants and algae only.
C. members of all three domains of life (bacteria, archaeans, eukaryotes). *
D. plants and bacteria only.
E. all bacteria, all archaeans, and certain eukaryotes.

There are organisms that carry out photosynthesis in all domains. [Stumbling Block 9]

7-8 CHAPTER 7 Copyright © 2013 by Nelson Education Ltd.

 5. C4 plants are most common in regions that are

A. hot, dry. *
B. temperate, dry.
C. hot, humid.
D. temperate but not dry.
E. cold.
F. There are no regions where C4 plants are common.

This question provides an opportunity for students to think about the conditions that would
lead to photorespiration (as C4 plants have a form of photosynthesis that avoids the issues
relating to photorespiration).

UNANSWERED QUESTIONS

Photosynthesis is considered by many to be the most important biological process on Earth.

In particular, directly or indirectly (through herbivorous animals), photosynthesis provides all
of our food requirements. Research on photosynthesis therefore is of high importance and is
likely to have significant benefit for humankind. For example, a complete understanding of
the chemistry of photosynthesis, the regulation of the process, and the genes that encode the
components of the process could be applicable to other endeavours of human interest, such
as solar energy conversion and the development of therapeutic drugs.
From research on agricultural crops, we have learned that photosynthesis is not a very
efficient process. Estimates are that only 1 to 2 percent of the solar energy that strikes the
planet’s surface is converted to new photosynthetic products. Research is being done to
learn enough about photosynthesis so that crop plants can be engineered to be more
efficient. An area of particular relevance here is photorespiration, which reduces the
efficiency of energy use in photosynthesis. It is hoped that research will give us a better
understanding of the biochemical control of photorespiration and provide clues about
breeding new, more energy-efficient plants.
Let us consider two specific avenues of research.
1. How is the efficiency of photosynthesis regulated?
The laboratory of David Kramer at Washington State University is interested in the
energetics and control of photosynthesis, the electron transfer reactions, the coupling of
electron transfer reactions to ATP synthesis, and photosynthesis in extreme environments.
As you have learned, energy conversion by the chloroplast involves the capture of light
energy and the channelling of that energy through an electron transfer system with the
eventual synthesis of NADPH and ATP.
At high concentrations, many of the intermediates produced in this energy conversion can
potentially destroy the photosynthetic apparatus, a phenomenon called photoinhibition. To
prevent such damage, the efficiency of some of the photosystem components is reduced by
the release of some of the energy as heat. Increased heat lowers the efficiency of

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-9

photosynthesis, however. Evidence from a range of studies indicates that the balance
between protection against photoinhibition and photosynthetic efficiency is important in
enabling plants to acclimate to environmental changes. Kramer’s group is doing research to
develop an understanding of the structure and function of ATP synthase and the
cytochrome complex and the effects of these components on the proton-motive force,
which is known to play a pivotal role in balancing photoinhibition and photosynthetic
efficiency. The results will illuminate how the specific mechanisms of photosynthesis
determine plant growth and survival. In addition, the technology developed as part of the
research may lead to applications in plant breeding and farming, providing farmers with a
means to assess the physiological states of the plants they are growing and, therefore, to
modify the conditions for optimal growth.
2. How are chloroplast thylakoid membrane–protein complexes assembled?
Research by Andrew Webber’s group at Arizona State University is directed at understanding
the formation of chloroplast thylakoid membrane–protein complexes. Those complexes are
key to the process of photosynthesis, but their assembly is not understood. Using molecular
biology and biochemistry techniques, Webber’s group is studying how the synthesis of
chloroplast proteins, some of which are encoded by genes in the chloroplast and others of
which are encoded by genes in the nucleus, is coordinated and regulated. The researchers are
also using molecular techniques to change specific amino acids in the chloroplast proteins with
the aim of elucidating how those amino acids are involved in the assembly and functioning of
the complexes. The results will add more detailed knowledge about the structure and function
of components that are key to the process of photosynthesis.

WHAT OTHER RESOURCES ARE AVAILABLE?

 The Virtual Cell Animation Collection (Molecular and Cellular Biology Learning
Center) has some useful photosynthesis animations:
http://vcell.ndsu.nodak.edu/animations/photosynthesis/index.htm

 An amusing YouTube video: Z scheme drill:

http://www.youtube.com/watch?v=XsZlPeT3D10

 The Photosynthesis Song on YouTube (cheesy, but your students might enjoy it):
http://www.youtube.com/watch?v=C1_uez5WX1o

 ScienceDaily. “Newly Identified Enzymes Help Plants Sense Elevated CO2 and
Could Lead to Water-Wise Crops”:
http://www.sciencedaily.com/releases/2009/12/091213164705.htm

 Gust, D., Moore, T.A., and Moore, A.L. 1994. Photosynthesis mimics as molecular
electronic devices. Engineering in Medicine and Biology Magazine, IEEE 13(1): 58–66. doi
10.1109/51.265775.

7-10 CHAPTER 7 Copyright © 2013 by Nelson Education Ltd.

  Natural Resources Canada. Does harvesting in Canada’s forests contribute to climate
change? http://cfs.nrcan.gc.ca/publications?id=26918

 AspenFACE (Free-Air Carbon Dioxide Enrichment) Experiment—a

multidisciplinary study to assess the effects of increasing tropospheric ozone and
carbon dioxide levels on the structure and function of northern forest ecosystems:
http://aspenface.mtu.edu/index.html

 While not directly focused on photosynthesis, there are some interesting aspects of
plants at this site, Plants-in-Motion:
http://plantsinmotion.bio.indiana.edu/plantmotion/starthere.html

 Recent study on the origin of eukaryotic photosynthesis: Price, D.C., Chan, C.X.,
Yoon, H.S., Yang, E.C., Qiu, H., Weber, A.P., Schwacke, R., Gross, J., Blouin, N.A.,
Lane, C., Reyes-Prieto, A., Durnford, D.G., Neilson, J.A., Lang, F.B., Gertraud
Burger, G., Steiner, J.M., Löffelhardt, W., Meuser, J.E., Posewitz, M.C., Ball, S.,
Arias, M.C., Henrissat, B., Coutinho, P.M., Rensing, S.A., Symeonidi, A.,
Doddapaneni, H., Green, B.R., Rajah, V.D., Boore, J., and Bhattacharya, D. 2012.
Cyanophora paradoxa genome elucidates origin of photosynthesis in algae and plants.
Science 335: 843–847.

Copyright © 2013 by Nelson Education Ltd. PHOTOSYNTHESIS 7-11