11.6 Plant sensory systems and responses  (Page 5/40)

Amyloplasts (also known as statoliths ) are specialized plastids that contain starch granules and settle downward in response to gravity. Amyloplasts are found in shoots and in specialized cells of the root cap. When a plant is tilted, the statoliths drop to the new bottom cell wall. A few hours later, the shoot or root will show growth in the new vertical direction.

The mechanism that mediates gravitropism is reasonably well understood. When amyloplasts settle to the bottom of the gravity-sensing cells in the root or shoot, they physically contact the endoplasmic reticulum (ER), causing the release of calcium ions from inside the ER. This calcium signaling in the cells causes polar transport of the plant hormone IAA to the bottom of the cell. In roots, a high concentration of IAA inhibits cell elongation. The effect slows growth on the lower side of the root, while cells develop normally on the upper side. IAA has the opposite effect in shoots, where a higher concentration at the lower side of the shoot stimulates cell expansion, causing the shoot to grow up. After the shoot or root begin to grow vertically, the amyloplasts return to their normal position. Other hypotheses—involving the entire cell in the gravitropism effect—have been proposed to explain why some mutants that lack amyloplasts may still exhibit a weak gravitropic response.

Growth responses

A plant’s sensory response to external stimuli relies on chemical messengers (hormones). Plant hormones affect all aspects of plant life, from flowering to fruit setting and maturation, and from phototropism to leaf fall. Potentially every cell in a plant can produce plant hormones. They can act in their cell of origin or be transported to other portions of the plant body, with many plant responses involving the synergistic or antagonistic interaction of two or more hormones. In contrast, animal hormones are produced in specific glands and transported to a distant site for action, and they act alone.

Plant hormones are a group of unrelated chemical substances that affect plant morphogenesis. Five major plant hormones are traditionally described: auxins (particularly IAA), cytokinins, gibberellins, ethylene, and abscisic acid. In addition, other nutrients and environmental conditions can be characterized as growth factors.

Auxins

The term auxin is derived from the Greek word auxein , which means "to grow." Auxins are the main hormones responsible for cell elongation in phototropism and gravitropism. They also control the differentiation of meristem into vascular tissue, and promote leaf development and arrangement. While many synthetic auxins are used as herbicides, IAA is the only naturally occurring auxin that shows physiological activity. Apical dominance—the inhibition of lateral bud formation—is triggered by auxins produced in the apical meristem. Flowering, fruit setting and ripening, and inhibition of abscission    (leaf falling) are other plant responses under the direct or indirect control of auxins. Auxins also act as a relay for the effects of the blue light and red/far-red responses.