Plant Tracheary Elements PDF

Plant Tracheary Elements
Hiroo Fukuda, University of Tokyo, Tokyo, Japan

Formation of tracheary elements, which are constituents of vessels and tracheids, is a model of cell differentiation and programmed cell death in plants.
Secondary article
Article Contents
. Introduction . Primary Xylem and Tracheary Elements . Cell Differentiation Pathway (Genes, Cell Death) . Zinnia as a Model System
Introduction
The xylem, which is a tissue specic to the vascular plants, is composed of tracheary elements (TEs), parenchyma cells and bres. TEs are characterized by the formation of secondary cell walls with annular, spiral, reticulate or pitted wall thickenings. At maturity, TEs lose their nuclei and cell contents, to leave a hollow tube that is part of a vessel or tracheid. Dierentiation into TEs from procambial initials in situ and from various types of cells in vitro is a model of cell dierentiation and programmed cell death in plants.
. Function in Seedling Growth
microtubules and actin laments and treatment with agents that depolymerize cytoskeletons indicate that there is a coordinated mechanism in which actin laments are involved in the reorganization of microtubules which, in turn, regulate the spatial disposition of secondary walls (Kobayashi et al., 1988). The dynamic change in microtubule organization is accompanied by an increase in the number of microtubules due to new expression of some of the tubulin genes.
Primary Xylem and Tracheary Elements

The primary xylem develops from procambial initials in situ. Procambial initials originate in the early embryo. In Arabidopsis, the procambial primordium, which consists of eight narrow cells, is formed by the late globular stage of the embryo, and it forms the primary xylem tissues in the torpedo-shaped embryo (Bowman, 1993). During vegetative growth, the apical meristem serves as a continuous source of the procambial initials. The dierentiation of TEs from the procambial initials is divided into four ontogenetic processes: cell origination; cell elongation; secondary wall deposition; and wall lysis and cell death. Xylem dierentiation can also be induced by wound stress and/or a combination of phytohormones in vitro. The continuity of the xylem along the plant axis may be a result of the steady polar ow of auxin from young leaves to roots (Sachs, 1981). Cytokinin from roots may also be a controlling factor in xylem dierentiation.
Deposition of wall materials

During the formation of secondary walls, levels of cellulose and hemicellulose increase and the deposition of pectin ceases, and a little later lignin deposition starts. Lignin and polysaccharides are deposited in association with each other. At the time of secondary wall formation, rosettes are located in the plasma membrane over regions of secondary wall thickenings, showing active synthesis of cellulose there. Xylan is a major component of the increased hemicellulose and its synthesis is due to an increase in xylan synthase activity during secondary wall formation. Several proteins, including an extensin-like protein, arabinogalactan proteins and glycine-rich protein 1.8 are specically associated with secondary walls of Tes (Fukuda, 1997). The biosynthesis of lignin involves three pathways, known as the shikimate, the general phenylpropanoid and the specic lignin pathways. The general phenylpropanoid pathway involves phenylalanine ammonia-lyase (PAL), cinnamate hydroxylase, O-methyltransferase, and 4-coumarate:CoA ligase (4CL), all of which are expressed in association with TE dierentiation. Promoters of genes encoding PAL and 4CL share conserved sequences, resulting in the coordinated expression of these genes in the xylem (Hatton et al., 1995). Hydroxycinnamoyl-CoA esters, which are nal products of the general phenylpropanoid pathway, are converted to their corresponding alcohols, which are polymerized into lignin on secondary walls. Genes encoding enzymes that catalyse these processes are also expressed coupled with TE dierentiation.
Cell Differentiation Pathway (Genes, Cell Death)

Pattern formation of secondary walls
Microtubules determine the wall pattern by dening the position and orientation of secondary walls, probably by guiding the movement of cellulose-synthesizing rosettes in the plasma membrane. Microtubule arrays change dramatically from a longitudinal to a transverse orientation prior to secondary wall formation in Zinnia. Double staining of
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net
Plant Tracheary Elements
Cell death
In dierentiating TEs, the visible degeneration of all organelles, including the nucleus, starts only after the collapse of the tonoplast, which occurs several hours after the secondary wall thickenings become visible. Following the collapse of the tonoplast, organelles such as Golgi bodies and the endoplasmic reticulum become swollen and then rupture. The nucleus is also degraded. This degradation is preceded by the tonoplast collapse but not by nuclear shrinkage and fragmentation, which are typical features of apoptosis. This implies that the disruption of the tonoplast is a critical step towards cell death in dierentiating TEs. Specic hydrolytic enzymes such as a 30 kDa cysteine protease, an RNase, and a 43 kDa S1-type nuclease are induced prior to the tonoplast collapse and seem to be accumulated in the vacuole (Fukuda, 1997). Because pH optima of these enzymes are around 5.5, these hydrolytic enzymes may function in the vacuole or in the cytoplasm after the collapse of the tonoplast. In addition, proteasomes in the cytoplasm may also be involved in cell death programme of TEs.
Function in Seedling Growth

The xylem plays a role in the transport of water to all the parts of the seedlings from the roots. TEs are emptied by the loss of all cell contents to form hollow tubes as a pathway for uids, which is totally dierent from the case of the blood vessel. Transpiration and root pressure are engines for water transport. Because water is sucked up by high negative pressure, TEs should have a reinforced cell wall to maintain the shape under such high pressure. The xylem is also important from applied and biotechnological perspectives, because biomaterials, such as cellulose and lignin in xylem, represent the predominant part of the terrestrial biomass and therefore play a signicant role in the carbon cycle.
References
Bowman J (1993) Arabidopsis. New York: Springer-Verlag. Demura T and Fukuda H (1994) Novel vascular cell-specic genes whose expression is regulated temporally and spatially during vascular system development. Plant Cell 6: 967981. Fukuda H (1997) Tracheary element dierentiation. Plant Cell 9: 1147 1156. Hatton D, Sablowski R, Yung M-H, Smith C, Schuch W and Bevan M (1995) Two classes of cis sequences contribute to tissue-specic expression of a PAL2 promoter in transgenic tobacco. Plant Journal 7: 859876. Kobayashi H, Fukuda H and Shibaoka H (1988) Interrelationship between the spatial disposition of actin laments and microtubules during the dierentiation of tracheary elements in cultured Zinnia cells. Protoplasma 143: 2937. Sachs T (1981) The control of the patterned dierentiation of vascular tissues. Advances in Botanical Research 9: 151262.
Zinnia as a Model System

Induction of TE dierentiation has been achieved in callus, in suspension-cultured cells, and in excised tissues and cells. Among the in vitro dierentiation systems, the most ecient is the system in which single mesophyll cells isolated from Zinnia leaves can dierentiate directly into TEs without intervening cell division (Fukuda, 1997). In the Zinnia system, initiation of dierentiation occurs by wounding and by a combination of auxin and cytokinin. Localized secondary wall thickenings begin synchronously at 60 h of culture, and cell death occurs a little later. The process of in vitro TE dierentiation is divided into three stages, I, II and III. Stage I, which immediately follows the induction of dierentiation, corresponds to the functional dedierentiation process. This stage occurs only in the case of in vitro dierentiation. In stage II, dedierentiated cells change to procambial cells, and then to TE precursor cells. During this stage, some vascular cell-specic genes such as TED2-TED4 are preferentially expressed (Demura and Fukuda, 1994). Final determination occurs in the TE precursor cells and they then enter stage III, involving TEspecic events such as secondary wall deposition and cell death. Endogenous brassinosteroids may be involved in the nal determination.
Further Reading
Aloni R (1987) Dierentiation of vascular tissues. Annual Review of Plant Physiology 38: 179204. Barnett JR (ed.) (1981) Xylem Cell Development. Tunbridge Wells, UK: Castle House. Fukuda H (1996) Xylogenesis: initiation, progression and cell death. Annual Review of Plant Physiology and Plant Molecular Biology 47: 299325. Roberts LW, Gahan PB and Aloni R (1988) Vascular Dierentiation and Plant Growth Regulators. Berlin: Springer-Verlag. Torrey JG, Fosket DE and Hepler PK (1971) Xylem formation: a paradigm of cytodierentiation in higher plants. American Scientist 59: 338352.
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Plant Tracheary Elements PDF

Uploaded by

Copyright:

Available Formats

Plant Tracheary Elements PDF

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Plant Tracheary Elements PDF

Uploaded by

Copyright:

Available Formats

Plant Tracheary Elements

Hiroo Fukuda, University of Tokyo, Tokyo, Japan

. Function in Seedling Growth

Primary Xylem and Tracheary Elements

Deposition of wall materials

Cell Differentiation Pathway (Genes, Cell Death)

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Plant Tracheary Elements

Function in Seedling Growth

Zinnia as a Model System

ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

You might also like