Xylem

Multiple cross sections of a flowering plant stem showing primary and secondary xylem and phloem^[1]

In vascular plants, xylem is one of the two types of transport tissue, phloem being the other. The word "xylem" is derived from classical Greek ξυλον (xylon), "wood", and indeed the best-known xylem tissue is wood, though it is found throughout the plant. Its basic function is to transport water but it also transports some nutrients through the plant.

Physiology of xylem

The xylem is responsible for the transport of 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 contain a number of organic chemicals as well. This transport is not powered by energy spent by the tracheary elements themselves, which are dead by maturity and no longer have living contents. Two phenomena cause xylem sap to flow:

• Transpirational pull: the most important cause of xylem sap flow is the evaporation of water from the surfaces of mesophyll cells to the atmosphere. This transpiration 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 stomata open 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^[2].

Anatomy of xylem

Xylem can be found:

• in vascular bundles, present in non-woody plants and non-woody parts of plants with wood
• 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.

Note that, 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 most distinctive cells found in xylem are the tracheary elements: tracheids and vessel elements. However, the xylem is a complex tissue of plants, which means that it includes more than one type of cell. In fact, xylem contains other kinds of cells, such as parenchyma, in addition to those that serve to transport water.

The branching pattern exhibited by xylem has been shown to follow Murray's law.^[3]

Primary and secondary xylem

Primary xylem is the xylem that is formed during primary growth from procambium. It includes protoxylem and metaxylem. Metaxylem develops after the protoxylem but before secondary xylem. It is distinguished by wider vessels and tracheids. As it develops, the xylem can become endarch or exarch.

Secondary xylem is the xylem that 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:

1. conifers (Coniferae): there are some six hundred 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 marketed as softwood.
2. angiosperms (Angiospermae): there are some quarter of a million to four hundred thousand species of angiosperms. Within this group secondary xylem has not been found in the monocots. In the remainder of the angiosperms, this secondary xylem may or may not be present; this may vary even within a species, depending on growing circumstances. In view of the size of this group, it will be no surprise that no absolutes apply to the structure of secondary xylem within the angiosperms. Many non-monocot angiosperms become trees, and the secondary xylem of these is marketed as hardwood.

Evolution of xylem

Photos showing xylem elements in the shoot of a fig tree (Ficus alba): crushed in hydrochloric acid, between slides and cover slips.

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 plants. Several groups of plants later developed pitted tracheid cells, it seems, through convergent evolution. In living plants, pitted tracheids do not appear in development until the maturation of the metaxylem (following the protoxylem).

In most plant, pitted tracheids function as the primary transport cells. The other type of tracheary element, besides the tracheid, is the vessel element. Vessel elements are joined by perforations into vessels. In vessels, water travels by bulk flow, as in a pipe, rather than by diffusion through cell membranes. The presence of vessels in xylem has been considered to be one of the key innovations that led to the success of the angiosperms^[4]. 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^[5]. 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.

See also

• Cohesion-tension theory
• Phloem
• Secondary growth
• Transpirational pull
• Vascular tissue
• Vascular bundle
• Tylosis

References

General references

• Campbell, Neil A.; Jane B. Reece (2002). Biology (6th ed.). Benjamin Cummings. ISBN 978-0805366242. 
• Kenrick, Paul; Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution Press. ISBN 1-56098-730-8. 
• Muhammad, A.F.; R. Sattler (1982). "Vessel Structure of Gnetum and the Origin of Angiosperms". American Journal of Botany (Botanical Society of America) 69 (6): 1004–21. doi:10.2307/2442898. http://links.jstor.org/sici?sici=0002-9122%28198207%2969%3A6%3C1004%3AVSOGAT%3E2.0.CO%3B2-P. 
• Melvin T. Tyree; Martin H. Zimmermann (2003). Xylem Structure and the Ascent of Sap (2nd ed.). Springer. ISBN 3-540-43354-6.  recent update of the classic book on xylem transport by the late Martin Zimmermann