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JOURNAL ARTICLE
Why Leaves Turn Red: Pigments called anthocyanins probably protect leaves from light damage by direct shielding and by scavenging free radicals
David W. Lee and Kevin S. Gould
American Scientist
Vol. 90, No. 6 (NOVEMBER-DECEMBER 2002), pp. 524-531 (8 pages)
Published by: Sigma Xi, The Scientific Research Honor Society
Many forests, like those spread throughout New England, have just changed color in a spectacular way, as they do each fall. The phenomenon is familiar as well as dramatic, yet why it should happen has been a longstanding enigma. When we were in school, the standard textbooks said that foliage changes color because the breakdown of green chlorophyll molecules unmasks other pigments, like the yellow-to-orange xanthophylls and the red-or-blue anthocyanins, which, we were told, serve no particular function during the autumn senescence of leaves. Now botanists know better.
Indeed, a completely new appreciation for these colorful pigments has developed over the past decade or so, in part from our studies of trees in the Harvard Forest, a nature sanctuary in central Massachusetts maintained for scientific research. There, during September and October, one sees the leaves on dozens of woody species changing color. In some plants, such as the witch hazel (Hamamelis virginiana), it is indeed the loss of chlorophyll that reveals yellow carotenoid pigments, just as the textbooks say. However, for the forest’s 70 percent of tree species that contain anthocyanin pigments (which produce colors ranging from brown to red, depending on how much chlorophyll the leaves retain), the story is quite different. For example, the brilliant fall foliage of the red oak (Quercus rubra) results from the accumulation of anthocyanin in the vacuoles (large, fluid-filled cavities) of cells lying just under the leaves’ upper epidermis layer.
Anthocyanins are elaborate pigment molecules, widespread among land plants. They account not only for the autumn hues of temperate woodlands, but also for the flushes of developing red foliage seen in tropical forests, on the undersurface of shaded leaves and in crop plants suffering drought or nutrient deficiency. But plants can also have other red pigments. Carotenoids, often rhodoxanthin, produce red color in the senescing leaves of some conifers as well as in the common box (Buxas sempervirens), which decorates many suburban lawns. Betalain pigments color leaves red in a single order of flowering plants, and a few other miscellaneous pigments produce burgundy hues in very rare cases. But of all the red pigments, the anthocyanins are the most widespread.
We have collaborated in studying anthocyanin pigments since 1993 and are beginning to develop some working hypotheses about their function. It’s curious that an understanding has been so long in coming, given the fact these red pigments have been subjected to scientific scrutiny for nearly 200 years.
The Discovery of Anthocyanins
1段:A和落叶边红有管。德国植物学家先提出,A保护秋天的植物的新陈代谢. 但限于当时技术水平,无法证实。后来有人做实验推测A可能吸收UV-B.
Anthocyanins had been observed for centuries as ”colored cell sap.” In 1835 the German botanist Ludwig Marquart gave them their name, deriving anthocyanin from the Greek anthos, meaning flower, and kyanos, meaning blue. Many long-standing misconceptions about anthocyanin function date from these early observations, notably that these pigments arise from the breakdown of chlorophyll during autumn.
Given how striking and attractive red foliage is, it may seem baffling that botanists remained ignorant about the phenomenon for so long. There are various reasons for this. First, because anthocyanins are responsible for the colors of fruits and flowers as well as of leaves, it was natural to concentrate on pigmentation in the former economically important organs, for which the function of anthocyanin seems obvious—to attract animals for pollination and seed dispersal. Second, because the discoveries of Richard Willstatter and his colleagues about the molecular structure of anthocyanins from 1912 to 1916 were made shortly after the rediscovery of Mendel’s laws of inheritance, the anthocyanins became an early subject of research in molecular genetics, rather than physiology. (Mendel’s peas had distinctively colored flowers because of anthocyanins.) Third, the discovery that light can in— duce anthocyanin production inspired molecular biologists to study how light exposure activates genes involved in anthocyanin synthesis, again at the expense of research into anthocyanin function.
Botanists of the late 19th-century, most notably the Germans who studied plant anatomy and physiology, noticed that anthocyanin production rises when a plant is subjected to low temperatures and high light conditions. This observation led to the popular explanations that anthocyanins protect the photosynthetic structures against intense sunlight and help to warm leaves by increasing their rates of metabolism. These scientists lacked the instrumentation and detailed knowledge of photosynthesis to test their ideas. In the mid-20th century, investigators became aware that ultraviolet (UV) radiation could induce anthocyanin synthesis, leading to the hypothesis that anthocyanins protect plant tissues against UV damage. But, as it turns out, anthocyanins absorb rather weakly in the UV-B region of the spectrum (wavelengths of 285—320 nanometers), which is most responsible for damage to biological tissues; other colorless flavonoid pigments that are equally, or more, abundant in the leaves absorb UV-B much more sh'ongly. Furthermore, anthocyanins are most commonly produced in the interiors of leaves and hence are poorly placed to protect leaves from the widespread effects of UV—B. These weaknesses were refuted by one of us (Lee) in 1987. So what good are anthocyanins to a leaf? Two recent discoveries have shed light on the mystery.
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