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Lukas Schreiber, University of Bonn, Bonn, Germany

The endodermis of plant roots is a single layer of cells separating the cortex from the central cylinder. Owing to characteristic chemical modifications (suberin and lignin) of radial endodermal cell walls (Casparian bands) it forms an apoplastic barrier influencing radial nutrient and water transport in roots.

Secondary article
Article Contents
. Introduction . Anatomy . Isolation and Chemical Composition . Function as Apoplastic Barrier . Development and Root Mutants

The central cylinder of roots is separated from the cortex by the endodermis, which is a tissue composed of a single layer of cells. Since its rst description by Caspary (1866), the root endodermis and its structure and function have been the object of much speculation. Because of the characteristic cell wall modications in the radial walls (Casparian bands) of the endodermis (Figure 1), it was argued that the endodermis should form a perfect apoplastic barrier for water and solvents radially diusing from the soil solution across the cortex to the central

cylinder of roots, where xylem vessels are located. However, recent results, based on investigation of the chemical composition and transport properties of endodermal cell walls, indicate that there can be a substantial apoplastic bypass across Casparian bands. In addition to its function as a transport barrier, the endodermal cell wall, and also the hypodermal cell wall, of roots forms an important barrier against invading microorganisms.

In root cross-sections a pronounced modication of the radial endodermal cell walls occurs that can be made visible by uorescence microscopy (Figure 1). Due to the deposition of phenolic compounds showing a strong autouorescence, endodermal cell walls are chemically dierent from normal plant cell walls, which are mainly composed of carbohydrates. Anatomically, three developmental states can be distinguished. In its primary developmental state, the endodermal cell wall is characterized by the deposition of Casparian bands on the radial cell walls. In many species, a secondary development state (deposition of a lipophilic suberin lamella on the inner surface of the primary cell wall) and a tertiary developmental state (deposition of U-shaped cell wall thickenings onto the radial and inner tangential cell walls of the endodermal cells) can be observed. Casparian bands as well as suberin lamellae have also been detected in the root hypodermis of many species (Wilson and Peterson, 1983). Furthermore, there are some indications that an endodermis with a Casparian band can also occur in stems and leaves (Lersten, 1997). The physiological function in stems and leaves, however, remained basically unknown until recently, when it was reported that the shoot endodermis is responsible for the perception of gravity (Fukaki et al., 1998).

Figure 1 C Section across a primary root of maize (Zea mays L.). Lignified and suberized cell walls (R, rhizodermal cell layer; H, hypodermal cell layer; E, endodermal cell layer; and X, xylem vessel) exhibit a bright autofluorescence (excitation light used: 395 440 nm). Bar, 100 mm. (Illustration: K. Hartmann and L. Schreiber.)

Isolation and Chemical Composition

If root sections are treated with a mixture of cell wall degrading enzymes (pectinases and cellulases), cell walls

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that are chemically dierent from normal carbohydrate walls will resist the enzymatic attack. Using this approach it has become possible to isolate highly puried samples of endodermal and hypodermal cell walls (Karahara and Shibaoka, 1992; Schreiber et al., 1994), which can be further used for the analysis of the chemical composition. After chemical degradation of isolated cell wall samples, the monomer composition of the biopolymers can be determined by modern analytical techniques such as gas chromatography and mass spectrometry. Endodermal cell walls as well as hypodermal cell walls were shown to be composed of a complex mixture of dierent compounds. Besides normal carbohydrates, further structural biopolymers such as suberin, lignin and proteins contribute signicantly to the dry mass of the isolated cell wall samples (Zeier and Schreiber, 1998; Zeier et al., 1999). Endodermal cell walls in their primary developmental state (Casparian bands) contained only traces of suberin and signicantly larger amounts of lignin, whereas endodermal cell walls in their secondary developmental state were characterized by a signicantly larger amount of the lipophilic biopolymer suberin. Finally, tertiary endodermal cell walls were basically characterized by the deposition of lignied cellulose walls onto the suberin lamella. The chemical composition of hypodermal root cell walls was also characterized by the occurrence of suberin and lignin.

on the radial hydraulic conductivity of roots indicated that the formation of an exodermis (root hypodermis with Casparian bands) can also contribute signicantly to the radial resistance of plant roots towards water transport (Zimmermann et al., 2000). However, there is also substantial evidence that the radial transport of organic molecules, such as the phytohormone abscisic acid, can have a signicant apoplastic component (Freundl et al., 1998). Thus, it must be concluded that apoplastic barriers in roots (endodermal and hypodermal cell walls) are by no means perfect barriers completely impermeable for water and dissolved ions and organic molecules. Depending on the developmental state and the dierentiation of endoand hypodermal cell walls and on the physico-chemical properties of the substances, properties of the apoplastic barrier in roots can vary and an apoplastic bypass may signicantly contribute to the radial transport in roots.

Development and Root Mutants

The genetics of root development and formation of the endodermis with Casparian bands has been analysed using Arabidopsis thaliana, which is a good model system due to its simple cellular organization. Initials deriving from the root meristem give rise to specic cell types resulting in the nal radial organization of plant roots. Analysis of cell lineages and pattern formation has indicated that cortical and endodermal cells are derived from the same initial. Dierent mutations leading to characteristic changes in the radial organization of roots have been characterized. In SHORTROOT mutants the endodermis is completely missing, whereas in SCARECROW mutants there is no segregation between endodermis and cortex cells (Scheres et al., 1995). It is postulated that the SHORTROOT gene controls the number of cell layers in the root cortex. Recently, shoot gravitropism mutants have been isolated, and it was found that the endodermis in shoots is responsible for the negative gravitropism of Arabidopsis thaliana (Fukaki et al., 1998). Most interestingly, the genes responsible for the shoot gravitropism have been found to be allelic to the shortroot and scarecrow genes involved in the formation of the root endodermis. Thus, the same genes seem to be involved in the radial pattern formation of roots as in shoots of Arabidopsis.

Function as Apoplastic Barrier

Endodermal as well as hypodermal cell walls have an important function as apoplastic barriers in the roots of higher plants. Due to the complicated mixture of four dierent biopolymers simultaneously occurring within the same cell wall, it can be postulated that these cell wall structures will form a signicant barrier to microorganisms. Cell wall degrading enzymes, which are normally secreted by pathogens trying to infect plant tissues, will not easily degrade endodermal cell walls. Radial transport of water and solutes across corn (Zea mays L.) roots has been measured using root pressure probes (Peterson et al., 1993; Steudle et al., 1993). Radial water ow across roots with primary endodermal cell walls (Casparian bands) was not increased when the endodermis was damaged by puncturing, indicating the endodermis did not represent a major barrier towards the ow of water. However, solute permeability of sodium chloride was signicantly increased when the root endodermis was punctured, which proved that the endodermis represented an important apoplastic barrier for the radial transport of solutes. This conrmed earlier works where it was also shown, using tracers and electron microscopy, that the endodermis formed an apoplastic barrier (Du Pont and Leonard, 1977; Robards and Robb, 1974). Recent results

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Fukaki H, Wysocka Diller J, Kato T et al. (1998) Genetic evidence that the endodermis is essential for shoot gravitropism in Arabidopsis thaliana. Plant Journal 14: 425430. Karahara I and Shibaoka H (1992) Isolation of Casparian strips from pea roots. Plant and Cell Physiology 33: 555561. Lersten NR (1997) Occurrence of endodermis with a Casparian strip in stem and leaf. The Botanical Review 63: 265272. Peterson CA, Murrmann M and Steudle E (1993) Location of the major barriers to water and ion movement in young roots of Zea mays L. Planta 190: 127136. Robards AW and Robb ME (1974) The entry of ions and molecules into roots: an investigation using electron-opaque tracers. Planta 120: 1 12. Scheres B, Di Laurenzio L, Willemsen V et al. (1995) Mutations aecting the radial organisation of the Arabidopsis root display specic defects throughout the embryonic axis. Development 121: 5362. Schreiber L, Breiner HW, Riederer M, Du ggelin M and Guggenheim R (1994) The Casparian strip of Clivia miniata Reg. roots: isolation, ne structure and chemical nature. Botanica Acta 107: 353361. Steudle E, Murrmann M and Peterson CA (1993) Transport of water and solutes across maize roots modied by puncturing the endodermis. Further evidence for the composite transport model of the root. Plant Physiology 103: 335349. Wilson CA and Peterson CA (1983) Chemical composition of the epidermal, hypodermal, endodermal and intervening cortical cell walls of various plant roots. Annals of Botany 51: 759769. Zeier J and Schreiber L (1998) Comparative investigation of primary and tertiary endodermal cell walls isolated from the roots of ve monocotyledoneous species: chemical composition in relation to ne structure. Planta 206: 349361.

Zeier J, Goll A, Yokoyama M, Karahara I and Schreiber L (1999) Structure and chemical composition of endodermal and rhizodermal hypodermal walls of several species. Plant, Cell and Environment 22: 271279. Zimmermann HM, Hartmann K, Schreiber L and Steudle E (2000) Chemical composition of apoplastic transport barriers in relation to radial hydraulic conductivity of corn roots (Zea mays L.). Planta 210: 302311.

Further Reading
Aeschbacher RA, Schiefelbein J and Benfey P (1994) The genetic and molecular basis of root development. Annual Review of Plant Physiology and Plant Molecular Biology 45: 2545. Rost TL and Bryant JA (1996) Root organization and gene expression patterns. Journal of Experimental Botany 47: 16131628. Scheres B and Wolkenfelt H (1998) The Arabidopsis root as a model to study plant development. Plant Physiology and Biochemistry 36: 21 32. Schreiber L, Hartmann K, Skrabs M and Zeier J (1999) Apoplastic barriers in roots: chemical composition of endodermal and hypodermal cell walls. Journal of Experimental Botany 50: 12671280. Steudle E and Frensch J (1996) Water transport in plants: role of the apoplast. Plant and Soil 187: 6779. Steudle E and Peterson CA (1998) How does water get through roots? Journal of Experimental Botany 49: 775788. Tasaka M, Kato T and Fukaki H (1999) The endodermis and shoot gravitropism. Trends in Plant Science 4: 103107.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group /