Anne Mie C. Emons

Cortical microtubules and microfibril deposition in the cell wall of root hairs ofEquisetum hyemale

SummaryThe cell wall of root hairs ofEquisetum hyemale is shown to be composed of three different cell wall textures. The growing cell wall at the tip of the hair is composed of a dispersed texture of microfibrils, which continues along the outside of the whole hair. With increasing distance from the tip an increasing number of helicoidally arranged lamellae is deposited. These findings correspond with the observed isotropism of young hairs in polarized light.Hairs of approximately 4 days old become positive birefringent, indicating that longitudinally oriented layers prevail over layers with a transverse direction. This phenomenon starts at the base of the hair. Full-grown hairs are positive birefringent up to the tip and concordantly show a thick additional inner cell wall layer which forms a helical pattern the length of the hair, with a mean microfibril angle of 25∘ with the cell axis.Cortical microtubules, subjacent to the dispersed, the helicoidal and the helical wall texture are axially aligned and, thus, not in coalignment with the last deposited microfibrils.Coated and smooth vesicles are present in the cortical cytoplasm of both growing and full-grown hairs. Electron-dense profiles (20 nm in diameter), surrounded by a halo (of 50 nm) were observed on the wall-plasmalemma interface in full-grown hairs only. A relation of these structures with microfibril deposition could not be demonstrated. They might represent channels transporting material to the wall, which, in full-grown hairs, is heavily impregnated with a tawny brown substance.The general hypothesis that cortical microtubule orientation directs microfibril deposition is disputed.

Microtubules Do Not Control Microfibril Orientation in a Helicoidal Cell Wall

SummaryThe secondary cell wall layer of the young root hair ofEquisetum hyemale (L) has a helicoidal texture. The cortical microtubules in these hairs maintain an axial alignment while microfibrils are being deposited with a different orientation in each subsequent layer. The role of cortical microtubules in microfibril orientation is disputed.

Plasma-membrane rosettes in root hairs of Equisetum hyemale

Particle arrangement in the plasma membrane during cell wall formation was investigated by means of the double-replica technique in root hairs of Equisetum hyemale. Particle density in the protoplasmic fracture face of the plasma membrane was higher than in the extraplasmic fracture face. Apart from randomly distributed particles, particle rosettes were visible in the PF face of the plasma membrane. The rosettes consisted of six particles arranged in a circle and had an outer diameter of approx. 26 nm. No gradient in the number of rosettes was found, which agrees with micrifibril deposition taking place over the whole hair. The particle rosettes were found individually, which might indicate that they spin out thin microfibrils as found in higher-plant cell walls. Indeed microfibril width in these walls, measured in shadowed preparations, is 8.5±1.5 nm. It is suggested that the rosettes are involved in microfibril synthesis. Non-turgid cells lacked microfibril imprints in the plasma membrane and no particle rosettes were present on their PF face. Fixation with glutaraldehyde caused, probably as a result of plasmolysis, the microfibril imprints to disappear together with the particle rosettes. The PF face of the plasma membrane of non-turgid hairs sometimes showed domains in which the intramembrane particles were aggregated in a hexagonal pattern. Microfibril orientation during deposition will be discussed.

Golgi Body Motility in the Plant Cell Cortex Correlates with Actin Cytoskeleton Organization

The actin cytoskeleton is involved in the transport and positioning of Golgi bodies, but the actin-based processes that determine the positioning and motility behavior of Golgi bodies are not well understood. In this work, we have studied the relationship between Golgi body motility behavior and actin organization in intercalary growing root epidermal cells during different developmental stages. We show that in these cells two distinct actin configurations are present, depending on the developmental stage. In small cells of the early root elongation zone, fine filamentous actin (F-actin) occupies the whole cell, including the cortex. In larger cells in the late elongation zone that have almost completed cell elongation, actin filament bundles are interspersed with areas containing this fine F-actin and areas without F-actin. Golgi bodies in areas with the fine F-actin exhibit a non-directional, wiggling type of motility. Golgi bodies in areas containing actin filament bundles move up to 7 μm s⁻¹. Since the motility of Golgi bodies changes when they enter an area with a different actin configuration, we conclude that the type of movement depends on the actin organization and not on the individual organelle. Our results show that the positioning of Golgi bodies depends on the local actin organization.

Helicoidal cell-wall texture in root hairs

It is shown that root hairs of most aquatic plants have a helicoidal cell-wall texture. Cell walls of root hairs of the aquatic/marshland plant Ranunculus lingua, however, have an axial microfibril alignment. The occurrence of a helicoidal wall texture is not limited to root hairs of aquatic plants: the terrestrial plant Zebrina purpusii has a helicoidal root-hair wall texture, too. With the exception of the grasses, the occurrence of root hairs with helicoidal cell walls pertains to species with predetermined root-hair-forming cells, trichoblasts. The rotation mode of the helicoid is species-specific. The average angle between fibrils of adjacent lamellae varies from 23° to 40°. In Hydrocharis morsus-ranae, cortical microtubules have a net-axial orientation and thus do not parallel nascent microfibrils. The deposition of the helicoidal cell wall is discussed.