ulticellular plants are complex organisms and their

orderly development requires an extraordinary measure

of coordination between cells. In order to coordinate
their activities, cells must be able to communicate with
each other. The principal means of intercellular communication within plants are the hormones. Hormones
are signal molecules that individually or cooperatively
direct the development of individual cells or carry information between cells and thus coordinate growth and
development. Plant hormones have been the subject of
intensive investigation since auxin was first discovered
almost a century ago.
The discussion of each hormone in this and subsequent chapters will begin with a review of biosynthesis
and metabolism. An understanding of hormone biochemistry makes it easier to understand what kinds of
molecules they are and how they may function. In addition, a lot of what is known about what these molecules
do and how they do it is based on studies of mutants
that interfere with their biosynthesis or metabolism.
The metabolic turnover of hormone molecules is also a
significant factor in the regulation of cellular activities.
This first of four chapters on plant hormones is
devoted to auxin. The following chapters will cover
gibberellins, cytokinins, abscisic acid, ethylene, and
brassinosteroids. In the case of each hormone, we will
address the same three basic questions: what is it, what
does it do, and how does it do it?
Because this is the first chapter on hormones, we
will begin with an introduction to the hormone concept
in plants. The balance of the chapter includes
• the biochemistry and metabolism of auxins,
• a review of auxin’s principal effects on growth and
development,
• how auxin controls cell enlargement,
• auxin transport in the plant, and
• auxin control of genetic expression.
18.1 THE HORMONE CONCEPT
IN PLANTS
The concept of hormones, the chemical messengers that
enable cells to communicate with one another, arose in
the study of mammalian physiology. The latter half
of the nineteenth century witnessed exciting advances
in physiology and medicine. By 1850, it was known
that blood-borne substances originating in the testis
conditioned sexual characteristics. At the same time,
physicians pursuing clinical studies had become interested in the effect of glandular extracts and secretions
on the course of various diseases. By the turn of the century, a number of substances that elicited specific effects
on the growth and physiology of mammals had been
305
306 Chapter 18 / Hormones I: Auxins
demonstrated and the concept that bodily functions
were coordinated by the production and circulation of
chemical substances was gaining wide acceptance. In
1905, the British physician E. H. Starling introduced
the term hormone (Gr., to excite or arouse) to describe
these chemical messengers.
Application of the hormone concept to plants may
be traced as far back as the observations of Duhamel
du Monceau in 1758. Du Monceau observed the formation of roots on the swellings that occur above girdle
wounds that interrupted the phloem tissues around the
stems of woody plants. In order to explain these and
similar phenomena, German botanist Julius Sachs (ca.
1860) postulated specific organ-forming substances in
plants. Sachs postulated that root-forming substances,
for example, produced in the leaves and migrating down
the stem, would account for the initiation of roots
above the wound. The real beginning of plant hormone
research, however, is found in a series of simple but
elegant experiments conducted by Charles Darwin (see
Box 18.1). It was Darwin’s observations and experiments that ultimately led F. W. Went, almost half a
century later, to describe a hormonal-like substance as
the causative agent when plants grew toward the light.
At about the same time, H. Fitting introduced the term
hormone into the plant physiology literature.
What are hormones? Hormones are naturally
occurring, organic molecules that, at low concentration, exert a profound influence on physiological
processes. In addition, hormones, as defined by animal
physiologists, are (1) synthesized in a discrete organ or
tissue, and (2) transported in the bloodstream to a specific
target tissue where they (3) control a physiological
response in a concentration-dependent manner. While
there are many parallels between animal and plant
hormones, there are also some significant differences.
Like animal hormones, plant hormones are naturally
occurring organic substances that profoundly influence
physiological processes at low concentration. The site
of synthesis and mode of transport for plant hormones,
however, is not always so clearly localized. Although
some tissues or parts of tissues may be characterized by
higher hormone levels than others, synthesis of plant
hormones appears to be much more diffuse and cannot
always be localized to discrete organs.
A hormone can serve effectively as a regulatory
signal only if the molecule has a limited lifetime within
the target cell. Any molecule sufficiently long-lived to be
used repeatedly would sacrifice its dynamic, regulatory
function. This means that the amount of a hormone in
a cellular pool must be closely regulated and exhibit a
rate of metabolic turnover that is rapid relative to the
response that it controls.
The amount of hormone available to a target cell
will be governed primarily by the rates at which active
hormone molecules enter (input) and exit (output)
the hormone pool. Hormones may enter the pool by
(1) de novo synthesis of the hormone, (2) retrieval of
active hormone from an inactive storage form, such
as a chemical conjugate, and (3) transport of hormone into the pool from a site elsewhere in the plant.
Principal means for removing hormone from the pool
once it has acted include: (1) oxidation or some other
form of chemical degradation that renders the molecule
inactive or (2) synthesis of an irreversibly deactivated
conjugate. Clearly, in order to understand the dynamic
regulation of hormone activity in plants, it is essential
to know something of these inputs and outputs. No
understanding of hormone function can be complete
without a working knowledge of hormone biosynthesis
and metabolism.
18.2 AUXIN IS DISTRIBUTED
THROUGHOUT THE PLANT
Auxin (fr. G. auxein, to increase) is the quintessential
plant hormone. Auxin was the first plant hormone to
be discovered and it has a principal role in the most
fundamental of plant responses—the enlargement of
plant cells. Auxin is synthesized in meristematic regions
and other actively growing organs such as coleoptile
Root
Seed
Coleoptile
0 0.5 1.0
Relative auxin activity
FIGURE 18.1 Auxin distribution in an oat seedling (Avena
sativa), showing higher concentrations of hormone in
the actively growing coleoptile and root apices. (Based
on data from Thimann, K. V. 1934. Journal of General
Physiology 18:23–34.)
18.3 The Principal Auxin in Plants is Indole-3-Acetic Acid (IAA) 307
NH
BOX 18.1
DISCOVERING
AUXIN
The experimental beginnings of plant hormone research
in general and auxins in particular can be traced to the
work of Charles Darwin. Although Darwin is best known
for his work on evolution, later in his career he developed
an interest in certain aspects of plant physiology. Some
of these studies were summarized in the book The Power
of Movement in Plants, co-authored by his son, Francis.
One of several ‘‘movements’’ studied by the Darwins
was the tendency of canary grass (Phalaris canariensis)
seedlings to bend toward the light coming from a window, a phenomenon we now know as phototropism.
The primary leaves of grass seedlings are enclosed in
a hollow, sheath-like structure, called the coleoptile,
which encloses and protects the leaves as they grow up
through the soil. Darwin observed that coleoptiles, like
stems, respond to unilateral illumination by growing
toward the light source. However, curvature would not
occur if the tip of the coleoptile were either removed
or covered in order to exclude light. Since the bending response was observed over the entire coleoptile,
Darwin concluded that the phototropic signal was perceived by the tip and ‘‘that when the seedlings are freely
exposed to lateral light, some influence is transmitted
from the upper to the lower part, causing the latter to
bend.’’ It was the implications of Darwin’s ‘‘transmissible influence’’ that captured the imagination of plant
physiologists and set into motion a series of experiments
that culminated in the discovery of the plant hormone,
auxin—the first plant hormone to be discovered.
Following the publication of Darwin’s book, a number of scientists confirmed and extended their observations.
In 1910, Boysen-Jensen demonstrated that the
stimulus would pass through an agar block and was
therefore chemical in nature. In 1918, Paal showed
that if the apex were removed and replaced asymmetrically, curvature would occur even in darkness. In the
climate of the time—Baylis and Starling’s characterization of animal hormones had appeared only a few
years earlier—plant physiologists were quick to interpret these observations as strong support for a plant
hormone.
The active substance was first successfully isolated
in 1928 by F. W. Went, then a graduate student working in his father’s laboratory in Holland. Following up
on the earlier work of Boysen-Jensen and Paal, Went
removed the apex of oat (Avena sativa) coleoptiles and
stood the apical pieces on small blocks of agar. Allowing
a period of time for the substance to diffuse from the
tissue into the agar block, he then placed each agar
block asymmetrically on a freshly decapitated coleoptile. The substance then diffused from the block into
the coleoptile, preferentially stimulating elongation of
the cells on the side of the coleoptile below the agar
block. Curvature of the coleoptile was due to differential cell elongation on the two sides. Moreover, the
curvature proved to be proportional to the amount of
active substance in the agar. Went’s work was particularly significant in two respects: first, he confirmed
the existence of regulatory substances in the coleoptile apex, and second, he developed a means for isolation
and quantitative analysis of the active substance. Because
Went used coleoptiles from Avena seedlings, his quantitative test became known as the Avena curvature test.
Substances active in this test were called auxin, from the
Greek auxein (to increase).
The results of Went’s studies naturally stimulated intensive efforts to isolate and identify the
active substance. One particularly active compound,
indole-3-acetic acid (IAA), was isolated from human
urine in 1934. This peculiar source was selected because
it