An immunofluorescence study of sectioned barley endosperm imaged by confocal laser scanning microscopy provided three-dimensional data on the relationship of microtubules to the cytoplasm, nuclei, and cell walls during development from 4 to 21 days after pollination (DAP). Microtubules play an important role throughout endosperm ontogeny. The syncytium is organized into units of nuclear-cytoplasmic domains by nuclear-based radial microtubule systems that appear to control the pattern of the first anticlinal walls at 5 to 6 DAP. After 7 DAP, phragmoplasts of two origins (interzonal and cytoplasmic) guide wall formation. Large compartments formed by the "free growing" walls in association with cytoplasmic phragmoplasts formed adventitiously at interfaces of opposing microtubule systems are subsequently subdivided by interzonal phragmoplastlcell plates to give rise to the starchy endosperm. During development of the aleurone layer from 8 to 21 DAP, the microtubule cycle is typical of plant histogenesis; cortical microtubules are hooplike, and preprophase bands of microtubules predict the division plane.
INTRODUCTIONIn spite of the enormous economic importance of cereal endosperm in agriculture and industry, surprisingly little is known about its development and evolution. The endosperm is an enigma in that (1) it is a product of double fertilization &e., the union of one sperm cell with the egg results in the diploid zygote and the other sperm with one or more nuclei results in the primary endosperm nucleus); (2) unlike the embryo, which follows generally recognized patterns of plant histogenesis, the endosperm is so unlike any other plant tissue that it has been called a monstrosity; and (3) the endosperm has proven extremely difficult to study because in its early stage, it is a delicate liquid coenocyte contained within the innermost chamber of the developing seed.The evolutionary introduction of the endosperm into the life cycle of angiosperms is unknown. Double fertilization occurs in gymnosperms (Friedman, 1992), but the second fertilization results in an additional embryo that competes for resources during development. In angiosperms, the second fertilization results in the primary endosperm nucleus from which develops the nutritive tissue. The endosperm could either be a delayed continuation of female gametophyte development triggered by the second fertilization event and therefore homologous to the nutritive gametophyte in gymnosperms (Thomas, 1907), or the endosperm may be a highly modified supernumerary embryo (Sargant, 1900). Support for the second hypothesis is amassed by Friedman (1994), who argues that the endosperm is a homolog of the embryo, or in other words, an organism in its ownTo whom correspondence should be addressed.right. Recent molecular data demonstrating the expression of the same or homologous genes in the embryo and endosperm (Aalen et al., 1994) lend support to this view. From this perspective, the endosperm emerges as a model system for the study of plant development. It is abundant...