Sunlight satisfies two very important needs of biological

organisms: energy and information. On the one hand,

radiant energy from the sun maintains the planet’s surface temperature in a
range suitable for life and, through
the process of photosynthesis, is the ultimate source of
energy that sustains most of life in our biosphere. Radiation, primarily in the form
of light, also provides critical
information about the environment—information that
is used by plants to regulate movement, trigger developmental events, and mark
the passage of time. The
importance of light in the life of green plants is reflected
in the study of photobiology, which encompasses not
only phenomena such as photosynthesis (which reflects
the role of sunlight as an energy source) but also
phenomena such as photomorphogenesis and photoperiodism, where sunlight
provides the necessary
information for proper plant development and the measurement of daylength,
respectively.
In order to fully appreciate the pervasive importance
of light to plants, it is necessary to understand something of the physical nature of
light and the molecules
with which light interacts in plants. In this chapter,
we will
• explore the physical nature of light and how light
interacts with matter,
• discuss some of the terminology used in describing
light and methods for measuring it,
• discuss briefly the characteristics of light in the
natural environment of plants, and
• review the principal pigments and pigment systems
found in plants.
The various ways in which light is used by plants to
power photosynthesis and regulate development will be
discussed throughout many of the subsequent chapters.

6.1 THE PHYSICAL NATURE

OF LIGHT
6.1.1 LIGHT IS ELECTROMAGNETIC
ENERGY, WHICH EXISTS
IN TWO FORMS
What is light? As Johnson recognized more than 200
years ago, ‘‘we all know what light is, but it is not easy
to tell what it is.’’ The simplest answer is that light
is a form of radiant energy, a narrow band of energy
within the continuous electromagnetic spectrum

93
94 Chapter 6 / The Dual Role of Sunlight: Energy and Information
= 400 500 600 700 800 nm

Cosmic rays Infrared

X rays Solar rays
Gamma rays Visible light
Ultra- Radio waves
violet Sound

Yello
Violet Blue Green Orange Red Infrared
w

10-14 10-12 10-10 10-8 10-6 10-4 10-2 1 102 104 106 cm

FIGURE 6.1 The electromagnetic spectrum. Visible radiation, or light, represents

only a very small portion of the total electromagnetic spectrum.

of radiation emitted by the sun (Figure 6.1). The term

‘‘light’’ describes that portion of the electromagnetic
spectrum that causes the physiological sensation of
vision in humans. In other words, light is defined by the
range of wavelengths—between 400 and approximately
700 nanometers—capable of stimulating the receptors
located in the retina of the human eye. Strictly speaking,
those regions of the spectrum we perceive as red, green,
or blue are called light, whereas the ultraviolet and
infrared regions of the spectrum, which our eyes cannot
detect (although they may have significant biological
effects), are referred to as ultraviolet or infrared
radiation, respectively. While the following discussion
will focus on light, it is understood that the principles
involved apply to radiant energy in the broader sense.
Like other forms of energy, light is a bit of an enigma
and is difficult to define. It is more easily described not by
what it is but by how it interacts with matter. Physicists
of the late nineteenth and early twentieth centuries
resolved that light has attributes of both continuous
waves and discrete particles. Both of these attributes are
important in understanding the biological role of light.

6.1.2 LIGHT CAN BE CHARACTERIZED

AS A WAVE PHENOMENON
The propagation of light through space is characterized
by regular and repetitive changes, or waves, in its electrical and magnetic
properties. Electromagnetic radiation
actually consists of two waves—one electrical and one
magnetic—that oscillate at 90◦ to each other and to the
direction of propagation (Figure 6.2). The wave properties of light may be
characterized by either wavelength
or frequency. The distance in space between wave
crests is known as the wavelength and is represented
by the Greek letter lambda (λ). Biologists commonly
express wavelengths in units of nanometers (nm), where
1 nm = 10-9 m. Frequency, represented by the Greek
letter nu (ν), is the number of wave crests, or cycles,
passing a point in space in one second. Frequency is thus
related to wavelength in the following way:
ν = c/λ (6.1)
where c is the speed of light (3 × 108 m s-1). Biologists
most commonly use wavelength to describe light and
other forms of radiation, although frequency is useful
in certain situations. Wavelengths of primary interest to photobiologists fall into
three distinct ranges:
ultraviolet, visible, and infrared (Table 6.1).

6.1.3 LIGHT CAN BE CHARACTERIZED

AS A STREAM OF DISCRETE
PARTICLES
When light is emitted from a source or interacts with
matter, it behaves as though its energy is divided into
discrete units or particles called photons. The energy
carried by a photon is called a quantum (pl. quanta), to
reflect the fact that the energy can be quantized, that is,
it can be divided into multiple units.
The energy carried by a photon (Eq) is related
to wavelength and frequency in accordance with the
following relationship:
E
q = hc/λ = hν (6.2)
Direction of
propagation
E
E
H
H
1 wavelength

FIGURE 6.2 Wave nature of light. Electric vectors (E) and
magnetic vectors (H) oscillate at 90◦ to each other.

6.1 The Physical Nature of Light 95

TABLE 6.1 Radiation of Principal Interest to Biologists.

Wavelength Average Energy
Color Range (nm) (kJ mol-1 photons)
Ultraviolet 100–400
UV-C 100–280 471
UV-B 280–320 399
UV-A 320–400 332
Visible 400–740
Violet 400–425 290
Blue 425–490 274
Green 490–550 230
Yellow 550–585 212
Orange 585–640 196
Red 640–700 181
Far-red 700–740 166
Infrared longer than 740 85
where h is a proportionality constant, called Planck’s
constant. The value of h is 6.62 × 10-34 J s photon-1.
Accordingly, the quantum energy of radiation is inversely
proportional to its wavelength or directly proportional to
its frequency. The symbol hν (pronounced ‘‘h nu’’) is
commonly used to represent a photon in figures and
diagrams.
Since both h and c are constants, the energy of
a photon is easily calculated for any wavelength of
interest. The following example illustrates a calculation
of the energy content of red light, with a representative
wavelength of 660 nm (6.6 × 10-7 m).
E
q = (6.62 × 10-34J s photon-1)
(3 × 108m s-1)/6.6 × 10-7m (6.3)
Solving for Eq:
E
q = 3.01 × 10-19J photon-1 (6.4)
For blue light, with a representative wavelength of
435 nm (4.35 × 10-7 m),
E
q = (6.62 × 10-34J s photon-1)
(3 × 108m s-1)/4.35 × 10-7m (6.5)
Again, solving for Eq:
E
q = 4.56 × 10-19J photon-1 (6.6)
As the above numbers indicate, the energy content
of a single photon is a very small number. However, the
Einstein-Stark law of photochemical equivalence states that
one photon can interact with only one electron. Thus,
in any irreversible photochemical reaction, the energy
of one photon may be used to convert one molecule of
reactant A to one molecule of product B.
A + hν → B (6.7)
Since one mole of any substance contains Avogadro’s number (N) of molecules (N =
6.023 × 1023
molecules mol-1), to convert one mole of reactant A
to one mole of product B would require N number of
photons. Thus, for practical purposes it is convenient
to multiply the energy of a single photon by Avogadro’s
number, which gives the value of energy for a mole of
photons. The energy carried by a mole of photons of red
light, for example, is 181,292 J mol-1, or 181 kJ mol-1
(Table 6.1). The energy carried by a mole of photons of
blue light is correspondingly 274 kJ mol-1. The concept
of a mole of photons is more useful than dealing with
individual photons. For example, as will become apparent in the following section,
the law of photochemical
equivalence states that a mole of photons of a particular wavelength would be
required to excite a mole of
pigment molecules.

6.1.4 LIGHT ENERGY CAN INTERACT

WITH MATTER
For light to be used by plants, it must first be absorbed.
The absorption of light by any molecule is a photophysical event involving internal
electronic transitions
(Figure 6.3A). The Gotthaus-Draper principle tells us
that only light that is absorbed can be active in a photochemical process. In
contrast to photophysical events,
photochemistry refers to any chemical reaction which
utilizes absorbed light to convert reactants to products,
that is, any light-dependent reaction (Equation 6.7).
Therefore, any photobiological phenomenon requires
the participation of a molecule that absorbs light. Such
a molecule may be defined as a pigment. Plants contain
a variety of pigments that are prominent visual features
and important physiological components of virtually
all plants. The characteristic green color of leaves, for
example, is due to a family of pigments known as the
chlorophylls. Chlorophyll absorbs the light energy used
in photosynthesis. The pleasing colors of floral petals

96 Chapter 6 / The Dual Role of Sunlight: Energy and Information

GROUND STATE
ENERGY LEVEL
Relative
Absorbance
(nm)
400
500
600
T1 700
F
P
21

S2
S1
S0
a. b.


FIGURE 6.3 The absorption light by a molecule. (A) An
energy level diagram depicting the various possible transitions when light is absorbed. A
nonexcited molecule
is said to be in the ground state (S0). Upon absorption
of light of wavelength of either λ1 or λ2, a molecule can
undergo an electronic transition (solid arrows) to a singlet excited state represented by
either S1 or S2, respectively. Within each singlet excited state exist various
internal energy states representing vibrational and rotational states (smaller horizontal
lines). Dashed arrows
represent radiationless decay through which energy is
given up primarily as heat. Fluorescence (F) is the emission of light from the lowest excited
singlet state. T1
represents the metastable excited triplet state. Energy
from the triplet excited state may be lost by radiationless
decay or by delayed emission of light known as phosphorescence (P). The triplet state is
sufficiently long-lived
to allow for photochemical reactions to occur. (B) An
absorption spectrum (solid line) is a graph in which
absorbance is plotted as a function of wavelength. Peaks
or absorption bands correspond to the principal excitation levels illustrated in the energy
diagram. Also shown
is a fluorescence emission spectrum (dashed line) that
corresponds to the emission of the absorbed energy as
light from the lowest excited singlet state (S1).

are due to the anthocyanin pigments that serve to attract

insects as pollen vectors. Other pigments, such as phytochrome, are present in
quantities too small to be visible
but nonetheless serve important roles in plant morphogenesis. These and other
important plant pigments will
be described later in this chapter.
What actually happens when a pigment molecule
absorbs light? Absorption of light by a pigment molecule
is a rapid, photophysical, electronic event, occurring
within a femtosecond (fs = 10-15 s). In accordance
with the First Law of Thermodynamics (Chapter 5),
the energy of the absorbed photon is transferred to an
electron in the pigment molecule during that extremely
short period of time. The energy of the electron is thus
elevated from a low energy level, the ground state, to a
higher energy level known as the excited, or singlet,
state. This change in energy level is illustrated graphically in Figure 6.3. Like
photons, the energy states
of electrons are also quantized, that is, an electron can
exist in only one of a series of discrete energy levels.
A photon can be absorbed only if its energy content
matches the energy re