In woody plants, a tylosis (plural: tyloses) is a bladder-like distension of a parenchyma cell into the lumen of adjacent vessels. The term tylosis summarises the physiological process and the resulting occlusion in the xylem of woody plants as response to injury or as protection from decay in heartwood. [1] It is a key process in wall one of the compartmentalization of decay in trees (CODIT) and other woody plants.
Observed in section under a microscope, tyloses appear as balloon-like protrusions emanating from axial paratracheal parenchyma cells into xylem vessels through pits linking the two. In some types, there may be a distinct barrier between the tyloses emanating from the pits into the vessels, while they may be barely distinguishable in other cases. [2]
Tyloses are outgrowths on parenchyma cells of xylem vessels of secondary heartwood. When the plant is stressed by drought or infection, tyloses will fall from the sides of the cells and "dam" up the vascular tissue to prevent further damage to the plant.
Tyloses can aid in the process of making sapwood into heartwood in some hardwood trees, especially in trees with larger vessels. [3] These blockages can be used in addition to gum plugs as soon as vessels become filled with air bubbles, and they help to form a stronger heartwood by slowing the progress of rot. [3]
Tylosis in the vessels of flowering plants counteracts the axial spread of fungal hyphae and other pathogens by slowing down their vertical spread with a physical barrier. A similar process occurs in gymnosperms, which block access to tracheids by closing the pits that join them to each other. [4]
The blocked vessels also provide a defense against the radial spread of pathogens, limiting their horizontal spread through the plant stem. Protection is stronger at the boundaries where annual rings meet.
The effectiveness of both vertical and horizontal barriers is affected by the speed at which they are established by tyloses, being typically faster in healthier plants. [4]
As a tree grows, its cambium adds an annual increment (or ring) of new wood, and in many species of trees the older wood towards the centre of the tree becomes less important for physiological processes like water and nutrient transportation, and this is converted by the tree to heartwood. This heartwood has no active defenses against infection, but is protected from infection and decay by the blockage of the xylem vessels with tyloses and various substances such as gums, resins and waxes containing high concentrations of volatile organic compounds such as terpenes that are toxic to insect larvae and tree pathogens such as bacteria and fungi. These products are produced by the cambium and transported to the centre of the stem by cellular structures called medullary rays radiating from the center of the stem [5] and then enter living axial paratracheal parenchyma cells. As the wood ages, the contents of the parenchyma cell burst into the dead vessel through the pit linking the two. The parenchyma cell then dies as its contents are disgorged into the empty space of the dry vessel and highly effective decay-inhibiting substances, notably tannin, are formed and absorbed by the adjacent vessel walls.
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Wood is a structural tissue found in the stems and roots of trees and other woody plants. It is an organic material – a natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin that resists compression. Wood is sometimes defined as only the secondary xylem in the stems of trees, or it is defined more broadly to include the same type of tissue elsewhere such as in the roots of trees or shrubs. In a living tree it performs a support function, enabling woody plants to grow large or to stand up by themselves. It also conveys water and nutrients between the leaves, other growing tissues, and the roots. Wood may also refer to other plant materials with comparable properties, and to material engineered from wood, woodchips, or fiber.
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of the 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.
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 embryonic origin that together carry out a specific function. Organs are then formed by the functional grouping together of multiple tissues.
Bark is the outermost layer of stems and roots of woody plants. Plants with bark include trees, woody vines, and shrubs. Bark refers to all the tissues outside the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner bark, which in older stems is living tissue, includes the innermost layer of the periderm. The outer bark on older stems includes the dead tissue on the surface of the stems, along with parts of the outermost periderm and all the tissues on the outer side of the periderm. The outer bark on trees which lies external to the living periderm is also called the rhytidome.
The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xylem inwards, towards the pith, and secondary phloem outwards, towards the bark.
In botany, the trunk is the stem and main wooden axis of a tree, which is an important feature in tree identification, and which often differs markedly from the bottom of the trunk to the top, depending on the species. The trunk is the most important part of the tree for timber production.
Lepidodendron is an extinct genus of primitive lycopodian vascular plants belonging the order Lepidodendrales. Like other Lepidodendrales, species of Lepidodendron grew as large-tree-like plants in wetland coal forest environments. They sometimes reached heights of 50 metres, and the trunks were often over 1 m (3.3 ft) in diameter. They are often known as "scale trees", due to their bark being covered in diamond shaped leaf-bases, from which leaves grew during earlier stages of growth. They thrived during the Carboniferous Period, and persisted until the end of the Permian around 252 million years ago. Sometimes erroneously called "giant club mosses", the genus was actually more closely related to modern quillworts than to modern club mosses. Within the form classification system used within paleobotany, Lepidodendron is both used for the whole plant as well as specifically the stems and leaves.
Oak wilt is a fungal disease caused by the organism Bretziella fagacearum that threatens Quercus spp. The disease is limited to the eastern half of the United States; first described in the 1940s in the Upper Mississippi River Valley. The pathogen penetrates xylem tissue, preventing water transport and causing disease symptoms. Symptoms generally consist of leaf discoloration, wilt, defoliation, and death. The disease is dispersed by insect vectors and to adjacent trees through underground root networks. However, human spread is the most consequential dispersal method. Moving firewood long distances can potentially transport diseases and invasive species.
Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue: the vascular cambium and the cork cambium. All the vascular tissues within a particular plant together constitute the vascular tissue system of that plant.
In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of stems and roots, causing them to elongate, and gives rise to primary tissue. Secondary growth occurs in most seed plants, but monocots usually lack secondary growth. If they do have secondary growth, it differs from the typical pattern of other seed plants.
Compartmentalization of decay in trees (CODIT) is a concept created by plant pathologist Alex Shigo after studying wood-decay fungus patterns.
Ophiognomonia clavigignenti-juglandacearum is a mitosporic fungus that causes the butternut canker, a lethal disease of butternut trees. It is also known to parasitize other members of the genus Juglans on occasion, and very rarely other related trees including hickories. The fungus is found throughout North America, occurring on up to 91% of butternut trees, and may be threatening the viability of butternut as a species.
Ophiostoma ulmi is a species of fungus in the family Ophiostomataceae. It is one of the causative agents of Dutch elm disease. It was first described under the name Graphium ulmi, and later transferred to the genus Ophiostoma.
A branch collar is the "shoulder" between the branch and trunk of woody plants; the inflammation formed at the base of the branch is caused by annually overlapping trunk tissue. The shape of the branch collar is due to two separate growth patterns, initially the branch grows basipetally, followed by seasonal trunk growth which envelops the branch.
Lepidodendrales or arborescent lycophytes are an extinct group of primitive, vascular, heterosporous, arborescent (tree-like) plants belonging to Lycopodiopsida. Members of Lepidodendrales are the best understood of the fossil lycopsids due to the vast diversity of Lepidodendrales specimens and the diversity in which they were preserved; the extensive distribution of Lepidodendrales specimens as well as their well-preservedness lends paleobotanists exceptionally detailed knowledge of the coal-swamp giants’ reproductive biology, vegetative development, and role in their paleoecosystem. The defining characteristics of the Lepidodendrales are their secondary xylem, extensive periderm development, three-zoned cortex, rootlike appendages known as stigmarian rootlets arranged in a spiralling pattern, and megasporangium each containing a single functional megaspore that germinates inside the sporangium. Many of these different plant organs have been assigned both generic and specific names as relatively few have been found organically attached to each other. Some specimens have been discovered which indicate heights of 40 and even 50 meters and diameters of over 2 meters at the base. The massive trunks of some species branched profusely, producing large crowns of leafy twigs; though some leaves were up to 1 meter long, most were much shorter, and when leaves dropped from branches their conspicuous leaf bases remained on the surface of branches. Strobili could be found at the tips of distal branches or in an area at the top of the main trunk. The underground organs of Lepidodendrales typically consisted of dichotomizing axes bearing helically arranged, lateral appendages serving an equivalent function to roots. Sometimes called "giant club mosses", they are believed to be more closely related to extant quillworts based on xylem, although fossil specimens of extinct Selaginellales from the Late Carboniferous also had secondary xylem.
A woody plant is a plant that produces wood as its structural tissue and thus has a hard stem. In cold climates, woody plants further survive winter or dry season above ground, as opposed to herbaceous plants that die back to the ground until spring.
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, photosynthesis takes place here, stores nutrients, and produces new living tissue. The stem can also be called halm or haulm or culms.
Laminated root rot also known as yellow ring rot is caused by the fungal pathogen Phellinus weirii. Laminated root rot is one of the most damaging root disease amongst conifers in northwestern America and true firs, Douglas fir, Mountain hemlock, and Western hemlock are highly susceptible to infection with P. weirii. A few species of plants such as Western white pine and Lodgepole pine are tolerant to the pathogen while Ponderosa pine is resistant to it. Only hardwoods are known to be immune to the pathogen.
Medullary rays, also known as vascular rays or pith rays, are cellular structures found in some species of wood. They appear as radial planar structures, perpendicular to the growth rings, which are visible to the naked eye. In a transverse section they appear as radiating lines from the centre of the log. In an axial section they may appear as a variety of transverse markings, depending on how close the section is to the plane of the ray. In a tangential section they may be hard to see at all.
Injury in plants is damage caused by other organisms or by the non-living (abiotic) environment to plants. Animals that commonly cause injury to plants include insects, mites, nematodes, and herbivorous mammals; damage may also be caused by plant pathogens including fungi, bacteria, and viruses. Abiotic factors that can damage plants include heat, freezing, flooding, lightning, ozone gas, and pollutant chemicals.
