Root pressure

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Primary and secondary roots in a cotton plant Primary and secondary cotton roots.jpg
Primary and secondary roots in a cotton plant

Root pressure is the transverse osmotic pressure within the cells of a root system that causes sap to rise through a plant stem to the leaves. [1]

Root pressure occurs in the xylem of some vascular plants when the soil moisture level is high either at night or when transpiration is low during the daytime. When transpiration is high, xylem sap is usually under tension, rather than under pressure, due to transpirational pull. At night in some plants, root pressure causes guttation or exudation of drops of xylem sap from the tips or edges of leaves. Root pressure is studied by removing the shoot of a plant near the soil level. Xylem sap will exude from the cut stem for hours or days due to root pressure. If a pressure gauge is attached to the cut stem, the root pressure can be measured.

Root pressure is caused by active distribution of mineral nutrient ions into the root xylem. Without transpiration to carry the ions up the stem, they accumulate in the root xylem and lower the water potential. Water then diffuses from the soil into the root xylem due to osmosis. Root pressure is caused by this accumulation of water in the xylem pushing on the rigid cells. Root pressure provides a force, which pushes water up the stem, but it is not enough to account for the movement of water to leaves at the top of the tallest trees. The maximum root pressure measured in some plants can raise water only to 6.87 meters, and the tallest trees are over 100 meters tall.

Role of endodermis

The endodermis in the root is important in the development of root pressure. The endodermis is a single layer of cells between the cortex and the pericycle. These cells allow water movement until it reaches the Casparian strip, made of suberin, a waterproof substance. The Casparian strip prevents mineral nutrient ions from moving passively through the endodermal cell walls. Water and ions move in these cell walls via the apoplast pathway. Ions outside the endodermis must be actively transported across an endodermal cell membrane to enter or exit the endodermis. Once inside the endodermis, the ions are in the symplast pathway. They cannot diffuse back out again but can move from cell to cell via plasmodesmata or be actively transported into the xylem. Once in the xylem vessels or tracheids, ions are again in the apoplast pathway. Xylem vessels and tracheids transport water up the plant but lack cell membranes. The Casparian strip substitutes for their lack of cell membranes and prevents accumulated ions from diffusing passively in apoplast pathway out of the endodermis. The ions accumulating interior to the endodermis in the xylem create a water potential gradient and by osmosis, water diffuses from the moist soil, across the cortex, through the endodermis and into the xylem.

Root pressure can transport water and dissolved mineral nutrients from roots through the xylem to the tops of relatively short plants when transpiration is low or zero. The maximum root pressure measured is about 0.6 megapascals but some species never generate any root pressure. The main contributor to the movement of water and mineral nutrients upward in vascular plants is considered to be the transpirational pull. However, sunflower plants grown in 100% relative humidity grew normally and accumulated the same amount of mineral nutrients as plants in normal humidity, which had a transpiration rate 10 to 15 times the plants in 100% humidity. [2] Thus, transpiration may not be as important in upward mineral nutrient transport in relatively short plants as often assumed.

Xylem vessels sometimes empty over winter. Root pressure may be important in refilling the xylem vessels. [3] However, in some species vessels refill without root pressure. [4]

Root pressure is often high in some deciduous trees before they leaf out. Transpiration is minimal without leaves, and organic solutes are being mobilized so decrease the xylem water potential. Sugar maple accumulates high concentrations of sugars in its xylem early in the spring, which is the source of maple sugar. Some trees "bleed" xylem sap profusely when their stems are pruned in late winter or early spring, e.g. maple and elm. Such bleeding is similar to root pressure only sugars, rather than ions, may lower the xylem water potential. In the unique case of maple trees, sap bleeding is caused by changes in stem pressure and not root pressure.

It is very likely that all grasses produce root pressure. In bamboos, root pressure is correlated with maximum height of a clone. [5]

Related Research Articles

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

Plant cells are the cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capability to perform photosynthesis and store starch, a large vacuole that regulates turgor pressure, the absence of flagella or centrioles, except in the gametes, and a unique method of cell division involving the formation of a cell plate or phragmoplast that separates the new daughter cells.

<span class="mw-page-title-main">Xylem</span> Water transport tissue in vascular plants

Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word xylem is derived from the Ancient Greek word ξύλον (xylon), meaning "wood"; the best-known xylem tissue is wood, though it is found throughout a plant. The term was introduced by Carl Nägeli in 1858.

<span class="mw-page-title-main">Vascular plant</span> Clade of land plants with xylem and phloem

Vascular plants, also called tracheophytes or collectively Tracheophyta, form a large group of land plants that have lignified tissues for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue to conduct products of photosynthesis. Vascular plants include the clubmosses, horsetails, ferns, gymnosperms, and angiosperms. Scientific names for the group include Tracheophyta, Tracheobionta and Equisetopsida sensu lato. Some early land plants had less developed vascular tissue; the term eutracheophyte has been used for all other vascular plants, including all living ones.

<span class="mw-page-title-main">Tissue (biology)</span> Group of cells having similar appearance and performing the same function

In biology, tissue is a historically derived biological organizational level between cells and a complete organ. A tissue is therefore often thought of as an assembly of similar cells and their extracellular matrix from the same origin that together carry out a specific function. Organs are then formed by the functional grouping together of multiple tissues.

<span class="mw-page-title-main">Plant nutrition</span> Study of the chemical elements and compounds necessary for normal plant life

Plant nutrition is the study of the chemical elements and compounds necessary for plant growth and reproduction, plant metabolism and their external supply. In its absence the plant is unable to complete a normal life cycle, or that the element is part of some essential plant constituent or metabolite. This is in accordance with Justus von Liebig’s law of the minimum. The total essential plant nutrients include seventeen different elements: carbon, oxygen and hydrogen which are absorbed from the air, whereas other nutrients including nitrogen are typically obtained from the soil.

<span class="mw-page-title-main">Endodermis</span> Inner layer of cortex in vascular plant roots

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 to restrict apoplastic flow of water to the inside. The endodermis is the boundary between the cortex and the stele.

<span class="mw-page-title-main">Sap</span> Fluid transported in xylem cells or phloem sieve tube elements of a plant

Sap is a fluid transported in xylem cells or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant.

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

Suberin, cutin and lignins are complex, higher plant epidermis and periderm cell-wall macromolecules, forming a protective barrier. Suberin, a complex polyester biopolymer, is lipophilic, and composed of long chain fatty acids called suberin acids, and glycerol. 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 and 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.

<span class="mw-page-title-main">Apoplast</span> Extracellular space, outside the cell membranes of plants

The apoplast is the extracellular space outside of plant cell membranes, especially the fluid-filled continuum of cell walls of adjacent cells where material can diffuse freely. Fluid and material flows occurring in any extracellular space are called apoplastic flow or apoplastic transport. The apoplastic pathway is one route by which water and solutes are transported and distributed to different places through tissues and organs, contrasting with the symplastic pathway.

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

The symplast of a plant is the region enclosed by the cell membranes, within which water and solutes can diffuse freely. By contrast the apoplast is any fluid-filled space within the cell wall and extracellular space. Neighbouring cells are interconnected by microscopic channels known as plasmodesmata that traverse the cell walls. These channels, allow the flow of small molecules such as sugars, amino acids, and ions between cells. Larger molecules, including transcription factors and plant viruses, can also be transported through with the help of actin structures. The symplast allows direct cytoplasm-to-cytoplasm flow of water and other nutrients along concentration gradients. In particular, symplastic flow is used in the root systems to bring in nutrients from soil. Nutrient solutes move in this way through three skin layers of the roots: from cells of the epidermis, the outermost layer, through the cortex into the endodermis.

<span class="mw-page-title-main">Vessel element</span> Component of Xylem

A vessel element or vessel member is one of the cell types found in xylem, the water conducting tissue of plants. Vessel elements are found in angiosperms but absent from gymnosperms such as conifers. Vessel elements are the main feature distinguishing the "hardwood" of angiosperms from the "softwood" of conifers.

A hydroid is a type of vascular cell that occurs in certain bryophytes. In some mosses such as members of the Polytrichaceae family, hydroids form the innermost layer of cells in the stem. At maturity they are long, colourless, thin walled cells of small diameter, containing water but no living protoplasm. Collectively, hydroids function as a conducting tissue, known as the hydrome, transporting water and minerals drawn from the soil. They are surrounded by bundles of living cells known as leptoids which carry sugars and other nutrients in solution. The hydroids are analogous to the tracheids of vascular plants but there is no lignin present in the cell walls to provide structural support.

The ascent of sap in the xylem tissue of plants is the upward movement of water and minerals from the root to the aerial parts of the plant. The conducting cells in xylem are typically non-living and include, in various groups of plants, vessel members and tracheids. Both of these cell types have thick, lignified secondary cell walls and are dead at maturity. Although several mechanisms have been proposed to explain how sap moves through the xylem, the cohesion-tension mechanism has the most support. Although cohesion-tension has received criticism due to the apparent existence of large negative pressures in some living plants, experimental and observational data favor this mechanism.

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

In plants, the transpiration stream is the uninterrupted stream of water and solutes which is taken up by the roots and transported via the xylem to the leaves where it evaporates into the air/apoplast-interface of the substomatal cavity. It is driven by capillary action and in some plants by root pressure. The main driving factor is the difference in water potential between the soil and the substomatal cavity caused by transpiration.

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. It was proposed by Ernst Münch, a German plant physiologist in 1930. A high concentration of organic substances, particularly sugar, inside cells of 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 known as hydrostatic pressure, in the phloem. Movement of phloem sap occurs by bulk flow from sugar sources to sugar sinks. The movement in phloem is bidirectional, whereas, in xylem cells, it is unidirectional (upward). Because of this multi-directional flow, coupled with the fact that sap cannot move with ease between adjacent sieve-tubes, it is not unusual for sap in adjacent sieve-tubes to be flowing in opposite directions.

<span class="mw-page-title-main">Plant stem</span> Structural axis of a vascular plant

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, stores nutrients, and produces new living tissue. The stem can also be called halm or haulm or culms.

<span class="mw-page-title-main">Transpiration</span> Process of water moving through a plant parts

Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. Water is necessary for plants but only a small amount of water taken up by the roots is used for growth and metabolism. The remaining 97–99.5% is lost by transpiration and guttation. Leaf surfaces are dotted with pores called stomata, and in most plants they are more numerous on the undersides of the foliage. The stomata are bordered by guard cells and their stomatal accessory cells that open and close the pore. Transpiration occurs through the stomatal apertures, and can be thought of as a necessary "cost" associated with the opening of the stomata to allow the diffusion of carbon dioxide gas from the air for photosynthesis. Transpiration also cools plants, changes osmotic pressure of cells, and enables mass flow of mineral nutrients and water from roots to shoots. Two major factors influence the rate of water flow from the soil to the roots: the hydraulic conductivity of the soil and the magnitude of the pressure gradient through the soil. Both of these factors influence the rate of bulk flow of water moving from the roots to the stomatal pores in the leaves via the xylem.

<span class="mw-page-title-main">Absorption of water</span> Life process in plants

In higher plants water and minerals are absorbed through root hairs which are in contact with soil water and from the root hairs zone a little the root tips.

<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. F. Baluska; Milada Ciamporová; Otília Gasparíková; Peter W. Barlow, eds. (2013). Structure and Function of Roots: Proceedings of the Fourth International Symposium on Structure and Function of Roots, June 20–26, 1993. Vol. 58 (illustrated ed.). Springer Science & Business Media. p. 195. ISBN   9789401731010.
  2. Tanner, W.; Beevers, H. (31 July 2001). "Transpiration, a prerequisite for long-distance transport of minerals in plants?". Proceedings of the National Academy of Sciences. 98 (16): 9443–9447. Bibcode:2001PNAS...98.9443T. doi: 10.1073/pnas.161279898 . PMC   55440 . PMID   11481499.
  3. Sperry, J. S.; Holbrook, N. M.; Zimmermann, M. H.; Tyree, M. T. (February 1987). "Spring Filling of Xylem Vessels in Wild Grapevine". Plant Physiology. 83 (2): 414–417. doi:10.1104/pp.83.2.414. PMC   1056371 . PMID   16665259.
  4. Tibbetts, T. J.; Ewers, F. W. (September 2000). "Root pressure and specific conductivity in temperate lianas: exotic Celastrus orbiculatus (Celastraceae) vs. native Vitis riparia (Vitaceae)". American Journal of Botany. 87 (9): 1272–1278. doi: 10.2307/2656720 . JSTOR   2656720. PMID   10991898.
  5. Cao, Kun-Fang; Yang, Shi-Jian; Zhang, Yong-Jiang; Brodribb, T.J. (July 2012). "Maximum height of grasses is determined by roots". Ecology Letters. 15 (7): 666–672. doi:10.1111/j.1461-0248.2012.01783.x. PMID   22489611.