Introduction

Have you hugged a tree lately? If not, you might want to give it some thought. You,
along with the rest of the human population, owe your existence to plants and other
organisms that capture light. In fact, most life on Earth is possible because the sun
provides a continuous supply of energy to ecosystems.

All organisms, including humans, need energy to fuel the metabolic reactions of growth,
development, and reproduction. But organisms can't use light energy directly for their
metabolic needs. Instead, it must first be converted into chemical energy through the
process of photosynthesis.

What is photosynthesis?

Photosynthesis is the process in which light energy is converted to chemical energy in the
form of sugars. In a process driven by light energy, glucose molecules (or other sugars)
are constructed from water and carbon dioxide, and oxygen is released as a byproduct.
The glucose molecules provide organisms with two crucial resources: energy and fixed—
organic—carbon.

 Energy. The glucose molecules serve as fuel for cells: their chemical energy can
be harvested through processes like cellular respiration and fermentation, which
generate adenosine triphosphate—ATP a small, energy-carrying molecule—for the
cell’s immediate energy needs.

 Fixed carbon. Carbon from carbon dioxide—inorganic carbon—can be

incorporated into organic molecules; this process is called carbon fixation, and the
carbon in organic molecules is also known as fixed carbon. The carbon that's fixed
and incorporated into sugars during photosynthesis can be used to build other
types of organic molecules needed by cells.
 In photosynthesis, solar energy is harvested and converted to chemical energy in the
form of glucose using water and carbon dioxide. Oxygen is released as a byproduct.

The ecological importance of photosynthesis

Photosynthetic organisms, including plants, algae, and some bacteria, play a key
ecological role. They introduce chemical energy and fixed carbon into ecosystems by
using light to synthesize sugars. Since these organisms produce their own food—that is,
fix their own carbon—using light energy, they are called photoautotrophs (literally, self-
feeders that use light).

Humans, and other organisms that can’t convert carbon dioxide to organic compounds
themselves, are called heterotrophs, meaning different feeders. Heterotrophs must get
fixed carbon by eating other organisms or their by-products. Animals, fungi, and many
prokaryotes and protists are heterotrophs.

Besides introducing fixed carbon and energy into ecosystems, photosynthesis also
affects the makeup of Earth’s atmosphere. Most photosynthetic organisms generate
oxygen gas as a byproduct, and the advent of photosynthesis—over 3.33 billion years
ago, in bacteria resembling modern cyanobacteria—forever changed life on Earth. These
bacteria gradually released oxygen into Earth’s oxygen-poor atmosphere, and the
increase in oxygen concentration is thought to have influenced the evolution of aerobic
life forms—organisms that use oxygen for cellular respiration. If it hadn’t been for those
ancient photosynthesizers, we, like many other species, wouldn't be here today!

Photosynthetic organisms also remove large quantities of carbon dioxide from the
atmosphere and use the carbon atoms to build organic molecules. Without Earth’s
abundance of plants and algae to continually suck up carbon dioxide, the gas would build
up in the atmosphere. Although photosynthetic organisms remove some of the carbon
dioxide produced by human activities, rising atmospheric levels are trapping heat and
causing the climate to change. Many scientists believe that preserving forests and other
expanses of vegetation is increasingly important to combat this rise in carbon dioxide
levels.

Leaves are sites of photosynthesis

Plants are the most common autotrophs in terrestrial—land—ecosystems. All green plant
tissues can photosynthesize, but in most plants, but the majority of photosynthesis
usually takes place in the leaves. The cells in a middle layer of leaf tissue called
the mesophyll are the primary site of photosynthesis.

Small pores called stomata—singular, stoma—are found on the surface of leaves in most

plants, and they let carbon dioxide diffuse into the mesophyll layer and oxygen diffuse
out.

Each mesophyll cell contains organelles called chloroplasts, which are specialized to carry
out the reactions of photosynthesis. Within each chloroplast, disc-like structures
called thylakoids are arranged in piles like stacks of pancakes that are known as grana—
singular, granum. The membrane of each thylakoid contains green-colored pigments
called chlorophylls that absorb light. The fluid-filled space around the grana is called
the stroma, and the space inside the thylakoid discs is known as the thylakoid space.
Different chemical reactions occur in the different parts of the chloroplast.
The light-dependent reactions and the Calvin cycle

Photosynthesis in the leaves of plants involves many steps, but it can be divided into two
stages: the light-dependent reactions and the Calvin cycle.

 The light-dependent reactions take place in the thylakoid membrane and require a

continuous supply of light energy. Chlorophylls absorb this light energy, which is
converted into chemical energy through the formation of two compounds, ATP
an energy storage molecule—and NADPH—a reduced (electron-bearing) electron
carrier. In this process, water molecules are also converted to oxygen gas—the
oxygen we breathe!
  The Calvin cycle, also called the light-independent reactions, takes place in the
stroma and does not directly require light. Instead, the Calvin cycle uses ATP
and NADPH text from the light-dependent reactions to fix carbon dioxide and
produce three-carbon sugars which join up to form glucose.

Schematic of the light-dependent reactions and Calvin cycle and how they're connected.

The light-dependent reactions take place in the thylakoid membrane. They require light,
and their net effect is to convert water molecules into oxygen, while producing ATP
molecules—from ADP and Pi—and NADPH molecules—via reduction of NADP+.

ATP and NADPH are produced on the stroma side of the thylakoid membrane, where
they can be used by the Calvin cycle.

The Calvin cycle takes place in the stroma and uses the ATP and NADPH from the light-
dependent reactions to fix carbon dioxide, producing three-carbon sugars—
glyceraldehyde-3-phosphate, or G3P, molecules.
The Calvin cycle converts ATP to ADP and Pi, and it converts NADPH to NADP+. The
ADP, Pi, and NADP+ can be reused as substrates in the light reactions.

Overall, the light-dependent reactions capture light energy and store it temporarily in
the chemical forms of ATP is broken down to release energy, and NADPH donates its
electrons to convert carbon dioxide molecules into sugars. In the end, the energy that
started out as light winds up trapped in the bonds of the sugars.

Cellular respiration is one of the most elegant, majestic, and fascinating

metabolic pathways on earth. At the same time, it’s also one of the
most complicated. When I learned about it for the first time, I felt like I
had tripped and fallen into a can of organic-chemistry-flavored
alphabet soup!

Luckily, cellular respiration is not so scary once you get to know it.
Let's start by looking at cellular respiration at a high level, walking
through the four major stages and tracing how they connect up to one
another.
Steps of cellular respiration

Overview of the steps of cellular respiration.

1. Glycolysis. Six-carbon glucose is converted into two pyruvates

(three carbons each). ATP and NADH are made. These reactions
take place in the cytosol.

2. Pyruvate oxidation. Pyruvate travels into the mitochondrial

matrix and is converted to a two-carbon molecule bound to
 coenzyme A, called acetyl CoA. Carbon dioxide is released and
NADH is made.

3. Citric acid cycle. The acetyl CoA combines with a four-carbon

molecule and goes through a cycle of reactions, ultimately
regenerating the four-carbon starting molecule. ATP (or, in some
cases, GTP), NADH, and FADH_2 are made, and carbon dioxide is
released. These reactions take place in the mitochondrial matrix.

4. Oxidative phosphorylation. The NADH and FADH_2 produced in

other steps deposit their electrons in the electron transport chain
in the inner mitochondrial membrane. As electrons move down
the chain, energy is released and used to pump protons out of the
matrix and into the intermembrane space, forming a gradient. The
protons flow back into the matrix through an enzyme called ATP
synthase, making ATP. At the end of the electron transport chain,
oxygen accepts electrons and takes up protons to form water.

During cellular respiration, a glucose molecule is gradually broken

down into carbon dioxide and water. Along the way, some ATP is
produced directly in the reactions that transform glucose. Much more
ATP, however, is produced later in a process called oxidative
phosphorylation. Oxidative phosphorylation is powered by the
movement of electrons through the electron transport chain, a series
of proteins embedded in the inner membrane of the mitochondrion.

These electrons come originally from glucose and are shuttled to the
electron transport chain by electron carriers NAD+NAD+start text, N,
A, D, end text, start superscript, plus, end
superscript and FADFADstart text, F, A, D, end text, which
become NADHNADHstart text, N, A, D, H, end text and FADH2FADH2
start text, F, A, D, H, end text, start subscript, 2, end subscript when
they gain electrons. To be clear, this is what's happening in the diagram
above when it says ++plus NADHNADHstart text, N, A, D, H, end
text or ++plus FADH2FADH2start text, F, A, D, H, end text, start
subscript, 2, end subscript. The molecule isn't appearing from scratch,
it's just being converted to its electron-carrying form:

NAD+NAD+start text, N, A, D, end text, start superscript, plus, end

superscript ++plus 2�−2e−2, e, start superscript, minus, end
superscript ++plus 2H+2H+2, start text, H, end text, start superscript,
plus, end superscript →→right arrow NADHNADHstart text, N, A, D,
H, end text ++plus H+H+start text, H, end text, start superscript, plus,
end superscript

FADFADstart text, F, A, D, end text ++plus 2�−2e−2, e, start

superscript, minus, end superscript ++plus 2H+2H+2, start text, H, end
text, start superscript, plus, end superscript →→right
arrow FADH2FADH2start text, F, A, D, H, end text, start subscript, 2,
end subscript

To see how a glucose molecule is converted into carbon dioxide and

how its energy is harvested as ATP and NADHNADHstart text, N, A, D,
H, end text//slashFADH2FADH2start text, F, A, D, H, end text, start
subscript, 2, end subscript in one of your body's cells, let’s walk step by
step through the four stages of cellular respiration.

1. Glycolysis. In glycolysis, glucose—a six-carbon sugar—undergoes

a series of chemical transformations. In the end, it gets converted
into two molecules of pyruvate, a three-carbon organic molecule.
In these reactions, ATP is made, and NAD+NAD+start text, N, A,
 D, end text, start superscript, plus, end superscript is converted
to NADHNADHstart text, N, A, D, H, end text.

2. Pyruvate oxidation. Each pyruvate from glycolysis goes into the

mitochondrial matrix—the innermost compartment of
mitochondria. There, it’s converted into a two-carbon molecule
bound to Coenzyme A, known as acetyl CoA. Carbon dioxide is
released and NADHNADHstart text, N, A, D, H, end text is
generated.

3. Citric acid cycle. The acetyl CoA made in the last step combines
with a four-carbon molecule and goes through a cycle of
reactions, ultimately regenerating the four-carbon starting
molecule. ATP, NADHNADHstart text, N, A, D, H, end text,
and FADH2FADH2start text, F, A, D, H, end text, start subscript,
2, end subscript are produced, and carbon dioxide is released.

4. Oxidative phosphorylation. The NADHNADHstart text, N, A, D, H,

end text and FADH2FADH2start text, F, A, D, H, end text, start
subscript, 2, end subscript made in other steps deposit their
electrons in the electron transport chain, turning back into their
"empty" forms (NAD+NAD+start text, N, A, D, end text, start
superscript, plus, end superscript and FADFADstart text, F, A, D,
end text). As electrons move down the chain, energy is released
and used to pump protons out of the matrix, forming a gradient.
Protons flow back into the matrix through an enzyme called ATP
synthase, making ATP. At the end of the electron transport chain,
oxygen accepts electrons and takes up protons to form water.

Glycolysis can take place without oxygen in a process

called fermentation. The other three stages of cellular respiration—
pyruvate oxidation, the citric acid cycle, and oxidative phosphorylation
—require oxygen in order to occur. Only oxidative phosphorylation
uses oxygen directly, but the other two stages can't run without
oxidative phosphorylation.

Each stage of cellular respiration is covered in more detail in other

articles and videos on the site. Try watching the overview video, or
jump straight to an article on a particular stage by using the links
above.