Electron microscopy of gas space (aerenchyma) formation in adventitious roots of Zea mays L. subjected to oxygen shortage

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Abstract
This paper examines the ultrastructure of cortical cells in maize root tips during the early stages in lysigenous aerenchyma formation, promoted by oxygen-deficient nutrient solution. The aim was to determine whether changes in fine structure were compatible with oxygen starvation as the primary cause of cell degeneration and death. There was an initial collapse of some cortical cells, indicating loss of turgor, and the cytoplasm became more electron dense. Mitochondria and endoplasmic reticulum appeared normal at this early stage though the tonoplast lost its integrity. Subsequently the cytoplasm became less electron dense than surrounding healthy cells, and underwent further degeneration while the plasmalemma retracted from the cell wall. Cell walls remained unaltered until this stage, but some then became thin and electron transparent. No cells of the stele were found to degenerate. These observations, which do not readily accord with the hypothesis that oxygen starvation was the cause of cell death, are compared with detailed studies of cell degeration in other cell types. An alternative mechanism for the stimulation of cortical cell lysis in poorly oxygenated roots involving the hormone ethylene, is discussed.