Ontogenesis of the collapsed layer during haustorium development in the root hemi-parasite Santalum album Linn.

The structure and development of collapsed layers of the haustorium were studied in Santalum album Linn. Through light and transmission electron microscopy, it was shown that the collapsed layers originated from starch-containing cells when the haustorium developed an internal gland, thickened gradually and ultimately developed into the mantle, which, combined with the sucker, buckled the host root. We report on the presence of inter-collapsed layers for the first time. These layers develop after penetration into the host and are located between the intrusive tissues and the vascular meristematic region, gradually linking the collapsed layers and remains around the sucker. The proliferation of cells in the meristematic region and the 'host tropism' of cortical layers contribute to pressure within the haustorium and result in development of the collapsed layers. Besides, starch-containing cells that turn into collapsed layers are vulnerable to pressure as they lack a large vacuole, have uneven cell wall thickness and a loose cell arrangement. We proposed that the functions of collapsed layers are to efficiently assure that cell inclusion and energy concentrate at the inner meristematic region and are recycled to affect penetration, reinforce the physical connection between the sandalwood haustorium and host root, and supply space for haustorial development.