INFLUENCE OF SHADE ON STOMATAL DENSITY, LEAF SIZE AND OTHER LEAF CHARACTERISTICS IN THE MAJOR TROPICAL ROOT CROPS, TANNIA, SWEET POTATO, YAM, CASSAVA AND TARO

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Abstract
Since shading often occurs under normal growing conditions in various cropping systems (for example, intercropping, alley cropping), field experiments were carried out to determine the effects of shading on stomatal density, leaf size, leaf dry matter, and leaf lamina thickness in the major tropical root and tuber crops, tannia (Xanthosoma sagittifolium), sweet potato (Ipomoea batatas), yam (Dioscorea esculenta), cassava (Manihot esculenta), and taro (Colocasia esculenta). Shading decreased stomatal density in the lower epidermis of tannia, sweet potato, yam and cassava, and in the upper epidermis also in tannia and sweet potato; the upper epidermis of yam and cassava were devoid of stomata. In contrast to the other species, taro under shade had an increased stomatal density in both the upper and lower epidermis, a finding which was confirmed in subsequent pot experiments. This response of taro was postulated as a possible manifestation of greater shade adaptation by the species.For all the species, shading generally resulted in the production of larger (in terms of surface area) but thinner leaves, with a decreased dry matter concentration. For the two species (yam and cassava) that had stomata only on the lower epidermis, normal sun-grown plants had about twice as many stomata per unit area of lower epidermis than did corresponding plants of the other three species. The agronomic and physiological significance of the findings are discussed.