Apoplast

Within a plant, the apoplast is the space outside the plasma membrane within which material can diffuse freely. It is interrupted by the Casparian strip in roots, by air spaces between plant cells and by the plant cuticle.

Structurally, the apoplast is formed by the continuum of cell walls of adjacent cells as well as the extracellular spaces, forming a tissue level compartment comparable to the symplast. The apoplastic route facilitates the transport of water and solutes across a tissue or organ. This process is known as apoplastic transport.

The apoplast is important for all the plant's interaction with its environment. The main carbon source (carbon dioxide) needs to be solubilized in the apoplast before it diffuses through the plasma membrane into the cell's cytoplasm (symplast) and is used by the chloroplasts during photosynthesis. In the roots, ions diffuse into the apoplast of the epidermis before diffusing into the symplast, or in some cases being taken up by specific ion channels, and being pulled by the plant's transpiration stream, which also occurs completely within the boundaries of the apoplast. Similarly, all gaseous molecules emitted and received by plants such as plant hormones and other pheromones must pass the apoplast. In nitrate poor soils, acidification of the apoplast increases cell wall extensibility and root growth rate. This is believed to be caused by a decrease in nitrate uptake (due to deficit in the soil medium) and supplanted with an increase in chloride uptake. H+ATPase increases the efflux of H+, thus acidifying the apoplast. The apoplast is also a site for cell-to-cell communication. During local oxidative stress, hydrogen peroxide and superoxide anions can diffuse through the apoplast and transport a warning signal to neighbouring cells. In addition, a local alkalinization of the apoplast due to such a stress can travel within minutes to the rest of the plant body via the xylem and trigger systemic acquired resistance. The apoplast also plays an important role in resistance to aluminum toxicity and resistance. Exclusion of aluminum ions in the apoplast prevent toxic levels which inhibit shoot growth, reducing crop yields

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