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The power of movement is not normally associated with

plants. Yet movement pervades the life of the green


plant. Movement in higher plants does not involve locomotion as it does in animals, nor is it so obvious. Plant
movement is mostly slow and deliberate, but it is a
key factor in determining the orientation of plants in
space. Plants that have been inadvertently placed in the
horizontal position will reorient their root and shoot
to the vertical. House plants will bend, appearing to
seek light coming through a window. Leaves may periodically rise and fall throughout the day and night,
while others track the sun as it moves across the sky.
Leaves of the Venus flytrap snap closed on a hapless
insect. While most plant movements are relatively slow,
they nonetheless serve important functions by positioning organs for the uptake of nutrients and water and
optimal interception of sunlight, or (in the case of the
flytrap) obtaining nutrients such as nitrogen through
the leaves.
There are two principal categories of movement in
plants, based on the distinctiveness of their mechanisms.
Growth movements are irreversible. They arise as
the result of differential growth within an organ or
between two different organs. Turgor movements
are reversible, resulting from simple volume changes in
certain cells—most often in a special organ called the
pulvinus. Within each group, we can further distinguish
between nutation, tropism, and nastic movement.
The term nutation (or circumnutations) denotes a
regular rotary or helical movement of plant organs, most
typically the stem apex, in space. Nutations are best demonstrated by time-lapse photography. Tropic responses
are directionally related to the stimulus such
as light (phototropism), gravity (gravitropism), water
(hydrotropism), or touch (thigmotropism). Nastic responses are not obviously related to any vector in the
stimulus. Directionality of nastic responses is inherent in the
tissue and includes epinasty (bending down), hyponasty
(bending up), nyctinasty (the rhythmic sleep movements
of leaves), seismonasty (response to mechanical shock),
thermonasty (temperature), and thigmonasty (touch).
This chapter will focus on the three plant movements that have been explored most thoroughly. These
are
• phototropism, particularly the nature of the photoreceptor and the role of auxin in the signal transduction
chain,
• gravitropism, including a brief discussion of the
nature of the gravitational stimulus and the mechanism of gravity perception, the particular character
of gravitropism in shoots and roots, and the role
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392 Chapter 23 / Tropisms and Nastic Movements: Orienting Plants in Space
of auxin and calcium in the differential growth response, and
• nastic movements, including the structure of motor
organs and the role of potassium flux in nyctinastic
and seismonastic movements.
23.1 PHOTOTROPISM:
REACHING FOR THE SUN
Most people are familiar with the sight of house plants
bending toward the light from an open window, an
everyday example of the phenomenon called phototropism (Figure 23.1). Phototropism is a classic plant
physiology problem that has attracted the interest of
botanists since the middle of the nineteenth century.
Darwin’s study of phototropism, published in his book
The Power of Movement in Plants in 1881, is credited
with overcoming the preoccupation of English-speaking
botanists with descriptive and taxonomic biology and
stimulating an interest in the more dynamic aspects of
plant function. Cell elongation in phototropically stimulated grass coleoptiles also led to Went’s discovery of
plant hormones (Chapter 18).
Tropic responses may be either positive or negative.
If a plant responds in the direction of the stimulus (e.g.,
toward a light source) it is said to be positive. If it
grows away from the stimulus it is said to be negative.
Whether the phototropic response is positive or negative
depends largely on the nature of the organ or its age.
For example, coleoptiles, hypocotyls, and the elongating
portions of stems and other aerial organs are for the
most part positively phototropic while the tendrils of
most climbing plants are negatively phototropic. Leaves
are normally plagiotropic, which means they orient at
FIGURE 23.1 The phototropic response in oat (Avena
sativa) coleoptiles. Left: Dark-grown seedling placed
in unilateral blue light, from the right, for 90 minutes.
Right: Unlighted control. Note that the overall length
of the coleoptile is approximately the same in the two
seedling