Endodermis

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Ranunculus Root Cross Section Ranunculus Root Cross Section.png
Ranunculus Root Cross Section

The endodermis is the innermost layer of cortex in land plants. It is a cylinder of compact living cells, the radial walls of which are impregnated with hydrophobic substances (Casparian strip) to restrict apoplastic flow of water to the inside. [1] The endodermis is the boundary between the cortex and the stele.

Contents

In many seedless plants, such as ferns, the endodermis is a distinct layer of cells immediately outside the vascular cylinder (stele) in roots and shoots. In most seed plants, especially woody types, the endodermis is present in roots but not in stems.

The endodermis helps regulate the movement of water, ions and hormones into and out of the vascular system. It may also store starch, be involved in perception of gravity and protect the plant against toxins moving into the vascular system.

Structure

The endodermis is developmentally the innermost portion of the cortex. It may consist of a single layer of barrel-shaped cells without any intercellular spaces or sometimes several cell layers. The cells of the endodermis typically have their primary cell walls thickened on four sides radial and transverse with suberin, a water-impermeable waxy substance which in young endodermal cells is deposited in distinctive bands called Casparian strips. These strips vary in width but are typically smaller than the cell wall on which they are deposited. If the endodermis is likened to a brick cylinder (e.g. a smokestack), with the bricks representing individual cells, the Casparian strips are analogous to the mortar between the bricks. In older endodermal cells, suberin may be more extensively deposited on all cell wall surfaces and the cells can become lignified, forming a complete waterproof layer.

Some plants have a large number of amyloplasts (starch containing organelles) in their endodermal cells, in which case the endodermis may be called a starch sheath.

Endodermis is often made visible with stains like phloroglucinol due to the phenolic and lipid nature of the Casparian strips or by the abundance of amyloplasts.

Function

The endodermis prevents water, and any solutes dissolved in the water, from passing through this layer via the apoplast pathway. Water can only pass through the endodermis by crossing the membrane of endodermal cells twice (once to enter and a second time to exit). Water moving into or out of the xylem, which is part of the apoplast, can thereby be regulated since it must enter the symplast in the endodermis. This allows the plant to control to some degree the movement of water and to selectively uptake or prevent the passage of ions or other molecules.

The endodermis does not allow gas bubbles to enter the xylem and helps prevent embolisms from occurring in the water column. [2]

Passage cells are endodermal cells of older roots which have retained thin walls and Casparian strips rather than becoming suberized and waterproof like the other cells around them, to continue to allow some symplastic flow to the inside. Experimental evidence suggests that passage cells function to allow transfer of solutes such as calcium and magnesium into the stele, in order to eventually reach the transpiration system. [3] For the most part, however, old roots seal themselves off at the endodermis, and only serve as a passageway for water and minerals taken up by younger roots "downstream".

Endodermal cells may contain starch granules in the form of amyloplasts. These may serve as food storage, and have been shown to be involved in gravitropism in some plants. [4]

See also

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<span class="mw-page-title-main">Plant cell</span> Type of eukaryotic cell present in green plants

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<span class="mw-page-title-main">Root</span> Basal organ of a vascular plant

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<span class="mw-page-title-main">Bark (botany)</span> Outermost layers of stems and roots of woody plants

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<span class="mw-page-title-main">Amyloplast</span> Type of plastid, double-enveloped organelles in plant cells

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<span class="mw-page-title-main">Suberin</span> Hydrophobic lipid polyester in plant cell walls

Suberin is a lipophilic, complex polyester biopolymer of plants, composed of long-chain fatty acids called suberin acids and glycerol. Suberin, interconnected with cutins and lignins, also complex macromolecules, form a protective barrier in the epidermal and peridermal cell walls of higher plants. Suberins and lignins are considered covalently linked to lipids and carbohydrates, respectively, and lignin is covalently linked to suberin, and to a lesser extent, to cutin. Suberin is a major constituent of cork, and is named after the cork oak, Quercus suber. Its main function is as a barrier to movement of water and solutes.

<span class="mw-page-title-main">Casparian strip</span> Thickening in the root endodermis of vascular plants

The Casparian strip is a band-like thickening in the center of the root endodermis of vascular plants. The composition of the region is mainly suberin, lignin and some structural proteins, which are capable of reducing the diffusive apoplastic flow of water and solutes into the stele and its width varies between species. The Casparian strip is impervious to water so can control the transportation of water and inorganic salts between the cortex and the vascular bundle, preventing water and inorganic salts from being transported to the stele through the apoplast, so that it must enter the cell membrane and move to the stele through the symplastic pathway, blocking the internal and external objects of the cell. The function of mass transportation are similar to that of animal tissues.. The development of the Casparian strip is regulated by transcription factors such as SHORT-ROOT (SHR), SCARECROW (SCR) and MYB36, as well as polypeptide hormone synthesised by midcolumn cells.

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<span class="mw-page-title-main">Symplast</span> Interconnected intracellular space of a plant

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<span class="mw-page-title-main">Gravitropism</span> Plant growth in reaction to gravity and bending of leaves and roots

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<span class="mw-page-title-main">Transpiration stream</span>

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The pressure flow hypothesis, also known as the mass flow hypothesis, is the best-supported theory to explain the movement of sap through the phloem of plants. It was proposed by Ernst Münch, a German plant physiologist in 1930. Organic molecules such as sugars, amino acids, certain hormones, and messenger RNAs are known to be transported in the phloem through the cells called sieve tube elements. According to the hypothesis, high concentration of organic substances, particularly sugar, inside the phloem at a source such as a leaf, creates a diffusion gradient that draws water into the cells from the adjacent xylem. This creates turgor pressure, also called hydrostatic pressure, in the phloem. The hypothesis states that this is why movement of sap in the plant flows from the sugar producers (sources) to sugar absorbers (sinks).

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A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, engages in photosynthesis, stores nutrients, and produces new living tissue. The stem can also be called the culm, halm, haulm, or thyrsus.

<span class="mw-page-title-main">Exodermis</span> Part of a plant

The exodermis is a physiological barrier that has a role in root function and protection. The exodermis is a membrane of variable permeability responsible for the radial flow of water, ions, and nutrients. It is the outer layer of a plant's cortex. The exodermis serves a double function as it can protect the root from invasion by foreign pathogens and ensures that the plant does not lose too much water through diffusion through the root system and can properly replenish its stores at an appropriate rate.

References

  1. Rost, Thomas L.; Michael Barbour; C. Ralph Stocking; Terence Murphy (2006). Plant Biology, 2nd Edition. Thompson, Brooks/Cole. ISBN   978-0-534-38061-8.
  2. Taiz and Zeiger (2015). Plant Physiology and Development. Sunderland, Massachusetts U.S.A: Sinauer Associates, Inc. p. 109. ISBN   978-1-60535-353-1.
  3. Peterson. C. A.; Enstone, D. E. (1996). "Functions of passage cells in the endodermis and exodermis of root". Physiologia Plantarum. 97 (3): 592–598. doi:10.1111/j.1399-3054.1996.tb00520.x.
  4. Involvement of the Vacuoles of the Endodermis in the Early Process of Shoot Gravitropism in Arabidopsis, Miyo Terao Morita1, Takehide Kato1, Kiyoshi Nagafusaa, Chieko Saitoc, Takashi Uedac, Akihiko Nakanoc and Masao Tasaka, 2002. The Plant Cell 14:47-56