SUMMARYHypericum species have translucent oil cavities; many species also have superficially similar black internal structures. Contradictory descriptions of both structures emerged from 19th century studies, which we review here. Our study of Hypericum perforatum L. leaves and petals shows typical schizogenous translucent oil cavities that develop with a uniseriate epithelium. Black structures, m contrast, are nodules, each composed ot several large cells lacking a central intercellular space and surrounded by a biseriate sheath of flattened cells. Nodule cells accumulate black, granular, but non-resinous contents. The 'streaks' that traverse petals are elongate tubular cavities or nodules, except for some chimerical tubes composed partly of each. Hypericum balearicum L. leaves have bulging, pustular cavities. A large cellular nodule forms first, by non-synchronous cell divisions within the leaf mesophyll, and is surrounded by a sheath of two to three flattened cell layers. The central cells then separate unevenly to form an irregular cavity into which isolated cells and clusters of cells intrude, some of which continue to divide as other cells degenerate. Little or no oil accumulates in the cavity, and the peripheral lining of cells is very irregular to flattened and empty. Both species studied exhibit secretory structures with previously undescribed anatomical features.
Neutral (storage) oil bodies occur in leaf mesophyll cells of many angiosperms, but their literature has been largely forgotten. We review this literature and provide a survey of 302 species and hybrids from mostly north-central US species representing 113 families. Freehand cross sections of fresh leaves stained with Sudan IV verified the presence of oil. In 71 species from 24 families we observed 1-15 oil bodies per mesophyll cell. The eudicot families Asteraceae, Caprifoliaceae, Lamiaceae, and Rosaceae had the highest number of species with oil bodies, whereas few or no species in the Apiaceae, Betulaceae, Fabaceae, and Scrophulariaceae had them. Only three of 19 monocot species sampled had oil bodies. Repeat sampling of a Malus (crabapple) cultivar and a Euonymus species showed conspicuous oil bodies in mid-summer and also in mid-autumn in both attached and recently shed leaves. Oil bodies in leaf mesophyll cells are conspicuous (visible in hand cross sections using moderate magnification in unstained water mounts) in numerous species, and they occur throughout the growing season in at least some species. Neutral oil bodies in leaf mesophyll cells are not mentioned in contemporary textbooks and advanced works, but they deserve recognition as significant cellular components of many taxa, in which they may be significant sources of commercial oils.
When buds form in summer or early fall, modified stipules act as bud scales and their adaxial epidermis secretes a resin that fills the bud. This secretory layer collapses in the dormant bud. Immature leaves, stipules, and leaf primordia occupy the center of the bud; all lack functional resin glands. In spring, stipules of emerging leaves develop an adaxial palisadelike secretory epidermis that becomes more ridged longitudinally in successive stipules. Marginal teeth of the first leaves to emerge are covered with trichomes and lack a secretory epidermis. In successive leaves the teeth become glandular and secrete resin as the lamina unrolls. Later in the season, marginal leaf glands account for much of the resin. Unspecialized hydathodes or extrafloral nectaries occur proximal to each glandular tip. Guttation of water or nectar occurs here through stomata located above a vein ending. On the basis of field observations and a laboratory feeding experiment, the resin seems to function mainly as an insect repellent. It may also reduce water loss from young leaves.
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