The attachment strategy of English ivy: a complex mechanism acting on several hierarchical levels.

English ivy (Hedera helix L.) is able to grow on vertical substrates such as trees, rocks and house plaster, thereby attaching so firmly to the surface that when removed by force typically whole pieces of the climbing substrate are torn off. The structural details of the attachment process are not yet entirely understood. We studied the attachment process of English ivy in detail and suggest a four-phase process to describe the attachment strategy: (i) initial physical contact, (ii) form closure of the root with the substrate, (iii) chemical adhesion, and (iv) shape changes of the root hairs and form-closure with the substrate. These four phases and their variations play an important role in the attachment to differently structured surfaces. We demonstrate that, in English ivy, different mechanisms work together to allow the plant's attachment to various climbing substrates and reveal the importance of micro-fibril orientation in the root hairs for the attachment based on structural changes at the subcellular level.