Xylem is one of the two types of transport tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts of the plants such as stems and leaves, but it also transports nutrients.
Etymology
The word xylem is derived from the Ancient Greek word (), 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. From p. 9: "Ich will die beiden welche von dem Cambium nach aussen und nach innen gebildet werden, Phloëm und Xylem nennen." (I will call the two parts of the permanent tissue, which are formed by the cambium outwardly and inwardly, "phloëm" and "xylem".)
Structure
The most distinctive xylem cells are the long tracheary elements that transport water. Tracheids and vessel elements are distinguished by their shape; vessel elements are shorter, and are connected together into long tubes that are called vessels.
Wood also contains two other type of cells: parenchyma and fibers.Xylem . Encyclopædia Britannica
Xylem can be found: in vascular bundles, present in non-woody plants and non-woody parts of woody plants in secondary xylem, laid down by a meristem called the vascular cambium in woody plants as part of a stelar arrangement not divided into bundles, as in many ferns.
In transitional stages of plants with secondary growth, the first two categories are not mutually exclusive, although usually a vascular bundle will contain primary xylem only.
The branching pattern exhibited by xylem follows Murray's law.
Primary and secondary xylem Primary xylem is formed during primary growth from procambium. It includes protoxylem and metaxylem. Metaxylem develops after the protoxylem but before secondary xylem. Metaxylem has wider vessels and tracheids than protoxylem.
Secondary xylem is formed during secondary growth from vascular cambium. Although secondary xylem is also found in members of the gymnosperm groups Gnetophyta and Ginkgophyta and to a lesser extent in members of the Cycadophyta, the two main groups in which secondary xylem can be found are: conifers (Coniferae): there are approximately 600 known species of conifers. All species have secondary xylem, which is relatively uniform in structure throughout this group. Many conifers become tall trees: the secondary xylem of such trees is used and marketed as softwood. angiosperms (Angiospermae): there are approximately 250,000 known species of angiosperms. Within this group secondary xylem is rare in the monocots. Many non-monocot angiosperms become trees, and the secondary xylem of these is used and marketed as hardwood.
Main function – upwards water transport The xylem, vessels and tracheids of the roots, stems and leaves are interconnected to form a continuous system of water-conducting channels reaching all parts of the plants. The system transports water and soluble mineral nutrients from the roots throughout the plant. It is also used to replace water lost during transpiration and photosynthesis. Xylem sap consists mainly of water and inorganic ions, although it can also contain a number of organic chemicals as well. The transport is passive, not powered by energy spent by the tracheary elements themselves, which are dead by maturity and no longer have living contents. Transporting sap upwards becomes more difficult as the height of a plant increases and upwards transport of water by xylem is considered to limit the maximum height of trees. Three phenomena cause xylem sap to flow: Pressure flow hypothesis: Sugars produced in the leaves and other green tissues are kept in the phloem system, creating a solute pressure differential versus the xylem system carrying a far lower load of solutes—water and minerals. The phloem pressure can rise to several MPa, far higher than atmospheric pressure. Selective interconnection between these systems allows this high solute concentration in the phloem to draw xylem fluid upwards by negative pressure. Transpirational pull: Similarly, the evaporation of water from the surfaces of mesophyll cells to the atmosphere also creates a negative pressure at the top of a plant. This causes millions of minute menisci to form in the mesophyll cell wall. The resulting surface tension causes a negative pressure or tension in the xylem that pulls the water from the roots and soil. Root pressure: If the water potential of the root cells is more negative than that of the soil, usually due to high concentrations of solute, water can move by osmosis into the root from the soil. This causes a positive pressure that forces sap up the xylem towards the leaves. In some circumstances, the sap will be forced from the leaf through a hydathode in a phenomenon known as guttation. Root pressure is highest in the morning before the opening of stomata and allow transpiration to begin. Different plant species can have different root pressures even in a similar environment; examples include up to 145 kPa in Vitis riparia but around zero in Celastrus orbiculatus.
The primary force that creates the capillary action movement of water upwards in plants is the adhesion between the water and the surface of the xylem conduits.Cruiziat, Pierre and Richter, Hanno. Plant Physiology . Sinauer Associates. Capillary action provides the force that establishes an equilibrium configuration, balancing gravity. When transpiration removes water at the top, the flow is needed to return to the equilibrium.
Transpirational pull results from the evaporation of water from the surfaces of cells in the leaves. This evaporation causes the surface of the water to recess into the pores of the cell wall. By capillary action, the water forms concave menisci inside the pores. The high surface tension of water pulls the concavity outwards, generating enough force to lift water as high as a hundred meters from ground level to a tree's highest branches.
Transpirational pull requires that the vessels transporting the water be very small in diameter; otherwise, cavitation would break the water column. And as water evaporates from leaves, more is drawn up through the plant to replace it. When the water pressure within the xylem reaches extreme levels due to low water input from the roots (if, for example, the soil is dry), then the gases come out of solution and form a bubble – an embolism forms, which will spread quickly to other adjacent cells, unless bordered pits are present (these have a plug-like structure called a torus, that seals off the opening between adjacent cells and stops the embolism from spreading). Even after an embolism has occurred, plants are able to refill the xylem and restore the functionality.
Cohesion-tension theory The cohesion-tension theory is a theory of intermolecular attraction that explains the process of water flow upwards (against the force of gravity) through the xylem of plants. It was proposed in 1894 by John Joly and Henry Horatio Dixon. Despite numerous objections, this is the most widely accepted theory for the transport of water through a plant's vascular system based on the classical research of Dixon-Joly (1894), Eugen Askenasy (1845–1903) (1895), and Dixon (1914,1924).
Water is a polar molecule. When two water molecules approach one another, the slightly negatively charged oxygen atom of one forms a hydrogen bond with a slightly positively charged hydrogen atom in the other. This attractive force, along with other intermolecular forces, is one of the principal factors responsible for the occurrence of surface tension in liquid water. It also allows plants to draw water from the root through the xylem to the leaf.
Water is constantly lost through transpiration from the leaf. When one water molecule is lost another is pulled along by the processes of cohesion and tension. Transpiration pull, utilizing capillary action and the inherent surface tension of water, is the primary mechanism of water movement in plants. However, it is not the only mechanism involved. Any use of water in leaves forces water to move into them.
Transpiration in leaves creates tension (differential pressure) in the cell walls of mesophyll cells. Because of this tension, water is being pulled up from the roots into the leaves, helped by cohesion (the pull between individual water molecules, due to hydrogen bonds) and adhesion (the stickiness between water molecules and the hydrophilic cell walls of plants). This mechanism of water flow works because of water potential (water flows from high to low potential), and the rules of simple diffusion.
Over the past century, there has been a great deal of research regarding the mechanism of xylem sap transport; today, most plant scientists continue to agree that the cohesion-tension theory best explains this process, but multiforce theories that hypothesize several alternative mechanisms have been suggested, including longitudinal cellular and xylem osmotic pressure gradients, axial potential gradients in the vessels, and gel- and gas-bubble-supported interfacial gradients.
Measurement of pressure
Until recently, the differential pressure (suction) of transpirational pull could only be measured indirectly, by applying external pressure with a pressure bomb to counteract it.The pressure of the water potential of the xylem in your plant's stem can be determined with the Scholander bomb. bio.usyd.edu.au When the technology to perform direct measurements with a pressure probe was developed, there was initially some doubt about whether the classic theory was correct, because some workers were unable to demonstrate negative pressures. More recent measurements do tend to validate the classic theory, for the most part. Xylem transport is driven by a combination of transpirational pull from above and root pressure from below, which makes the interpretation of measurements more complicated.
Evolution Xylem appeared early in the history of terrestrial plant life. Fossil plants with anatomically preserved xylem are known from the Silurian (more than 400 million years ago), and trace fossils resembling individual xylem cells may be found in earlier Ordovician rocks. The earliest true and recognizable xylem consists of tracheids with a helical-annular reinforcing layer added to the cell wall. This is the only type of xylem found in the earliest vascular plants, and this type of cell continues to be found in the protoxylem (first-formed xylem) of all living groups of vascular plants. Several groups of plants later developed pitted tracheid cells independently through convergent evolution. In living plants, pitted tracheids do not appear in development until the maturation of the metaxylem (following the protoxylem).
In most plants, pitted tracheids function as the primary transport cells. The other type of vascular element, found in angiosperms, is the vessel element. Vessel elements are joined end to end to form vessels in which water flows unimpeded, as in a pipe. The presence of xylem vessels (also called trachea) is considered to be one of the key innovations that led to the success of the angiosperms. However, the occurrence of vessel elements is not restricted to angiosperms, and they are absent in some archaic or "basal" lineages of the angiosperms: (e.g., Amborellaceae, Tetracentraceae, Trochodendraceae, and Winteraceae), and their secondary xylem is described by Arthur Cronquist as "primitively vesselless". Cronquist considered the vessels of Gnetum to be convergent with those of angiosperms. Whether the absence of vessels in basal angiosperms is a primitive condition is contested, the alternative hypothesis states that vessel elements originated in a precursor to the angiosperms and were subsequently lost.