e 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 391 392 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. L
76ftsdvnfiZZZZZZZZZZZzzvshidfrthuj Iorklgfvijlsdfahudnvhfyufyuvngyersgzdhjzderhuzfhfhjbfhjbvhund Thought Leaders That Have A Sincere Interest in Implementing Fundamental Changes That Not Only