Bark (botany)

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The bark of Pinus thunbergii is made up of countless shiny layers. Hei Song Pinus thunbergii 20211007185113 01.jpg
The bark of Pinus thunbergii is made up of countless shiny layers.

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. [1] 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. [2] [3] [4] [5] [6] [7] [8] [9] [10] [11]

Contents

Products derived from bark include bark shingle siding and wall coverings, spices, and other flavorings, tanbark for tannin, resin, latex, medicines, poisons, various hallucinogenic chemicals, and cork. Bark has been used to make cloth, canoes, and ropes and used as a surface for paintings and map making. [12] A number of plants are also grown for their attractive or interesting bark colorations and surface textures or their bark is used as landscape mulch. [13] [14]

The process of removing bark is decortication and a log or trunk from which bark has been removed is said to be decorticated. [15] [16] [17] [18] [19]

Botanical description

Bark is present only on woody plants - herbaceous plants and stems of young plants lack bark.

Tree cross section diagram Tree secondary components diagram.png
Tree cross section diagram

From the outside to the inside of a mature woody stem, the layers include the following: [20]

  1. Bark
    1. Periderm
      1. Cork (phellem or suber), includes the rhytidome
      2. Cork cambium (phellogen)
      3. Phelloderm
    2. Cortex
    3. Phloem
  2. Vascular cambium
  3. Wood (xylem)
    1. Sapwood (alburnum)
    2. Heartwood (duramen)
  4. Pith (medulla)

In young stems, which lack what is commonly called bark, the tissues are, from the outside to the inside:

  1. Epidermis, which may be replaced by periderm
  2. Cortex
  3. Primary and secondary phloem
  4. Vascular cambium
  5. Secondary and primary xylem.

Cork cell walls contain suberin, a waxy substance which protects the stem against water loss, the invasion of insects into the stem, and prevents infections by bacteria and fungal spores. [21] The cambium tissues, i.e., the cork cambium and the vascular cambium, are the only parts of a woody stem where cell division occurs; undifferentiated cells in the vascular cambium divide rapidly to produce secondary xylem to the inside and secondary phloem to the outside. Phloem is a nutrient-conducting tissue composed of sieve tubes or sieve cells mixed with parenchyma and fibers. The cortex is the primary tissue of stems and roots. In stems the cortex is between the epidermis layer and the phloem, in roots the inner layer is not phloem but the pericycle. [22] [23] [24] [25] [26] [27] [28] [29] [3]

As the stem ages and grows, changes occur that transform the surface of the stem into the bark. The epidermis is a layer of cells that cover the plant body, including the stems, leaves, flowers and fruits, that protects the plant from the outside world. In old stems the epidermal layer, cortex, and primary phloem become separated from the inner tissues by thicker formations of cork. Due to the thickening cork layer these cells die because they do not receive water and nutrients. This dead layer is the rough corky bark that forms around tree trunks and other stems.

Cork, sometimes confused with bark in colloquial speech, is the outermost layer of a woody stem, derived from the cork cambium. It serves as protection against damage from parasites, herbivorous animals and diseases, as well as dehydration and fire.

Periderm

Damaged bark of a cherry tree Poskozena borka na tresni 20210821 150436.jpg
Damaged bark of a cherry tree

Often a secondary covering called the periderm forms on small woody stems and many non-woody plants, which is composed of cork (phellem), the cork cambium (phellogen), and the phelloderm. The periderm forms from the phellogen which serves as a lateral meristem. The periderm replaces the epidermis, and acts as a protective covering like the epidermis. Mature phellem cells have suberin in their walls to protect the stem from desiccation and pathogen attack. Older phellem cells are dead, as is the case with woody stems. The skin on the potato tuber (which is an underground stem) constitutes the cork of the periderm. [30] [31]

In woody plants, the epidermis of newly grown stems is replaced by the periderm later in the year. As the stems grow a layer of cells form under the epidermis, called the cork cambium, these cells produce cork cells that turn into cork. A limited number of cell layers may form interior to the cork cambium, called the phelloderm. As the stem grows, the cork cambium produces new layers of cork which are impermeable to gases and water and the cells outside the periderm, namely the epidermis, cortex and older secondary phloem die. [32]

Within the periderm are lenticels, which form during the production of the first periderm layer. Since there are living cells within the cambium layers that need to exchange gases during metabolism, these lenticels, because they have numerous intercellular spaces, allow gaseous exchange with the outside atmosphere. As the bark develops, new lenticels are formed within the cracks of the cork layers.

Rhytidome

The rhytidome is the most familiar part of bark, being the outer layer that covers the trunks of trees. It is composed mostly of dead cells and is produced by the formation of multiple layers of suberized periderm, cortical and phloem tissue. [33] The rhytidome is especially well developed in older stems and roots of trees. In shrubs, older bark is quickly exfoliated and thick rhytidome accumulates. [34] It is generally thickest and most distinctive at the trunk or bole (the area from the ground to where the main branching starts) of the tree.

Chemical composition

Bark tissues make up by weight between 10 and 20% of woody vascular plants and consists of various biopolymers, tannins, lignin, suberin and polysaccharides. [35] Up to 40% of the bark tissue is made of lignin, which forms an important part of a plant, providing structural support by crosslinking between different polysaccharides, such as cellulose. [35]

Condensed tannin, which is in fairly high concentration in bark tissue, is thought to inhibit decomposition. [35] It could be due to this factor that the degradation of lignin is far less pronounced in bark tissue than it is in wood. It has been proposed that, in the cork layer (the phellogen), suberin acts as a barrier to microbial degradation and so protects the internal structure of the plant. [35] [36]

Analysis of the lignin in the bark wall during decay by the white-rot fungi Lentinula edodes (Shiitake mushroom) using 13C NMR revealed that the lignin polymers contained more Guaiacyl lignin units than Syringyl units compared to the interior of the plant. [35] Guaiacyl units are less susceptible to degradation as, compared to syringyl, they contain fewer aryl-aryl bonds, can form a condensed lignin structure, and have a lower redox potential. [37] This could mean that the concentration and type of lignin units could provide additional resistance to fungal decay for plants protected by bark. [35]

Damage and repair

Bark can sustain damage from environmental factors, such as frost crack and sun scald, as well as biological factors, such as woodpecker and boring beetle attacks. Male deer and other male members of the Cervidae (deer family) can cause extensive bark damage during the rutting season by rubbing their antlers against the tree to remove their velvet.

Living tree bark enveloping barbed wire BarkAndWire-0308.jpg
Living tree bark enveloping barbed wire

The bark is often damaged by being bound to stakes or wrapped with wires. In the past, this damage was called bark-galling and was treated by applying clay laid on the galled place and binding it up with hay. [38] In modern usage, "galling" most typically refers to a type of abnormal growth on a plant caused by insects or pathogens.

Bark damage can have several detrimental effects on the plant. Bark serves as a physical barrier to disease pressure, especially from fungi, so its removal makes the plant more susceptible to disease. Damage or destruction of the phloem impedes the transport of photosynthetic products throughout the plant; in extreme cases, when a band of phloem all the way around the stem is removed, the plant will usually quickly die. Bark damage in horticultural applications, as in gardening and public landscaping, results in often unwanted aesthetic damage.

The degree to which woody plants are able to repair gross physical damage to their bark is quite variable across species and type of damage. Some are able to produce a callus growth which heals over the wound rapidly, but leaves a clear scar, whilst others such as oaks do not produce an extensive callus repair. Sap is sometimes produced to seal the damaged area against disease and insect intrusion.[ citation needed ]

A number of living organisms live in or on bark, including insects, [39] fungi and other plants like mosses, algae and other vascular plants. Many of these organisms are pathogens or parasites but some also have symbiotic relationships.

Bark of mature mango (Mangifera indica) showing lichen growth Mango Bark.jpg
Bark of mature mango (Mangifera indica) showing lichen growth

Uses

The inner bark (phloem) of some trees is edible. In hunter-gatherer societies and in times of famine, it is harvested and used as a food source. In Scandinavia, bark bread is made from rye to which the toasted and ground innermost layer of bark of scots pine or birch is added. The Sami people of far northern Europe use large sheets of Pinus sylvestris bark that are removed in the spring, prepared and stored for use as a staple food resource. The inner bark is eaten fresh, dried or roasted. [40]

Bark of pine was used as emergency food in Finland during famine, last time during and after civil war in 1918. Detaching inner bark of pine.jpg
Bark of pine was used as emergency food in Finland during famine, last time during and after civil war in 1918.

Bark can be used as a construction material, and was used widely in pre-industrial societies. Some barks, particularly Birch bark, can be removed in long sheets and other mechanically cohesive structures, allowing the bark to be used in the construction of canoes, as the drainage layer in roofs, for shoes, backpacks, and other useful items. [41] Bark was also used as a construction material in settler colonial societies, particularly Australia, both as exterior wall cladding and as a roofing material. [42] [43]

Backpack made of birch bark. Museum by Lake Baikal, Russia Tragekorb aus Birkenleder.JPG
Backpack made of birch bark. Museum by Lake Baikal, Russia

In the cork oak ( Quercus suber ) the bark is thick enough to be harvested as a cork product without killing the tree; [44] in this species the bark may get very thick (e.g. more than 20 cm has been reported [45] ).

Some stem barks have significantly different phytochemical content from other parts. Some of these phytochemicals have pesticidal, culinary, or medicinally and culturally important ethnopharmacological properties. [46]

Bark contains strong fibres known as bast, and there is a long tradition in northern Europe of using bark from coppiced young branches of the small-leaved lime ( Tilia cordata ) to produce cordage and rope, used for example in the rigging of Viking Age longships. [47]

Among the commercial products made from bark are cork, cinnamon, quinine [48] (from the bark of Cinchona) [49] and aspirin (from the bark of willow trees). The bark of some trees, notably oak (Quercus robur) is a source of tannic acid, which is used in tanning. Bark chips generated as a by-product of lumber production are often used in bark mulch. Bark is important to the horticultural industry since in shredded form it is used for plants that do not thrive in ordinary soil, such as epiphytes. [50]

Bark chips Rindenmulch016.JPG
Bark chips

Wood bark contains lignin which when pyrolyzed yields a liquid bio-oil product rich in natural phenol derivatives. These are used as a replacement for fossil-based phenols in phenol-formaldehyde (PF) resins used in Oriented Strand Board (OSB) and plywood. [51]

See also

Related Research Articles

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

In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the surface of the soil, but roots can also be aerial or aerating, that is, growing up above the ground or especially above water.

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

Phloem is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as photosynthates, in particular the sugar sucrose, to the rest of the plant. This transport process is called translocation. In trees, the phloem is the innermost layer of the bark, hence the name, derived from the Ancient Greek word φλοιός (phloiós), meaning "bark". The term was introduced by Carl Nägeli in 1858. Different types of phloem can be distinguished. The early phloem formed in the growth apices is called protophloem. Protophloem eventually becomes obliterated once it connects to the durable phloem in mature organs, the metaphloem. Further, secondary phloem is formed during the thickening of stem structures.

<span class="mw-page-title-main">Tissue (biology)</span> Group of similar cells performing a specific function

In biology, tissue is an assembly of similar cells and their extracellular matrix from the same embryonic origin that together carry out a specific function. Tissues occupy a biological organizational level between cells and a complete organ. Accordingly, organs are formed by the functional grouping together of multiple tissues.

<span class="mw-page-title-main">Vascular cambium</span> Main growth tissue in the stems, roots of plants

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.

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

Cork cambium is a tissue found in many vascular plants as a part of the epidermis. It is one of the many layers of bark, between the cork and primary phloem. The cork cambium is a lateral meristem and is responsible for secondary growth that replaces the epidermis in roots and stems. It is found in woody and many herbaceous dicots, gymnosperms and some monocots. It is one of the plant's meristems – the series of tissues consisting of embryonic disk cells from which the plant grows. The function of cork cambium is to produce the cork, a tough protective material.

<span class="mw-page-title-main">Meristem</span> Type of plant tissue involved in cell proliferation

The meristem is a type of tissue found in plants. It consists of undifferentiated cells capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells continue to divide until they become differentiated and lose the ability to divide.

<span class="mw-page-title-main">Lenticel</span> Tissue that allows gas exchange in plant organs

A lenticel is a porous tissue consisting of cells with large intercellular spaces in the periderm of the secondarily thickened organs and the bark of woody stems and roots of gymnosperms and dicotyledonous flowering plants. It functions as a pore, providing a pathway for the direct exchange of gases between the internal tissues and atmosphere through the bark, which is otherwise impermeable to gases. The name lenticel, pronounced with an, derives from its lenticular (lens-like) shape. The shape of lenticels is one of the characteristics used for tree identification.

<span class="mw-page-title-main">Trunk (botany)</span> Main wooden axis of a tree

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.

<i>Lepidodendron</i> Extinct genus of vascular plants of the Carboniferous to Triassic

Lepidodendron is an extinct genus of primitive lycopodian vascular plants belonging the order Lepidodendrales. It is well preserved and common in the fossil record. 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 having been covered in diamond shaped leaf-bases, from which leaves grew during earlier stages of growth. However, they are correctly defined as arborescent lycophytes. 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. In the form classification system used in paleobotany, Lepidodendron is both used for the whole plant as well as specifically the stems and leaves.

<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">Cortex (botany)</span> Outer layer of a stem or root in a vascular plant

In botany, a cortex is an outer layer of a stem or root in a vascular plant, lying below the epidermis but outside of the vascular bundles. The cortex is composed mostly of large thin-walled parenchyma cells of the ground tissue system and shows little to no structural differentiation. The outer cortical cells often acquire irregularly thickened cell walls, and are called collenchyma cells.

<span class="mw-page-title-main">Ground tissue</span> Category of tissue in plants

The ground tissue of plants includes all tissues that are neither dermal nor vascular. It can be divided into three types based on the nature of the cell walls. This tissue system is present between the dermal tissue and forms the main bulk of the plant body.

  1. Parenchyma cells have thin primary walls and usually remain alive after they become mature. Parenchyma forms the "filler" tissue in the soft parts of plants, and is usually present in cortex, pericycle, pith, and medullary rays in primary stem and root.
  2. Collenchyma cells have thin primary walls with some areas of secondary thickening. Collenchyma provides extra mechanical and structural support, particularly in regions of new growth.
  3. Sclerenchyma cells have thick lignified secondary walls and often die when mature. Sclerenchyma provides the main structural support to the plant.
<span class="mw-page-title-main">Epidermis (botany)</span> Layer of cells that covers leaves, flowers, roots of plants

The epidermis is a single layer of cells that covers the leaves, flowers, roots and stems of plants. It forms a boundary between the plant and the external environment. The epidermis serves several functions: it protects against water loss, regulates gas exchange, secretes metabolic compounds, and absorbs water and mineral nutrients. The epidermis of most leaves shows dorsoventral anatomy: the upper (adaxial) and lower (abaxial) surfaces have somewhat different construction and may serve different functions. Woody stems and some other stem structures such as potato tubers produce a secondary covering called the periderm that replaces the epidermis as the protective covering.

<span class="mw-page-title-main">Vascular tissue</span> Conducting tissue in vascular plants

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.

<span class="mw-page-title-main">Secondary growth</span> Type of growth in plants

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.

The unifacial cambium produces cells to the interior of its cylinder. These cells differentiate into xylem tissue. Unlike the more common bifacial cambium found in later woody plants, the unifacial cambium does not produce phloem to its exterior. Also in contrast to the bifacial cambium, the unifacial cambium is unable to expand its circumference with anticlinal cell division. Cell elongation provides a limited amount of expansion.

<span class="mw-page-title-main">Lepidodendrales</span> Extinct order of vascular tree-like plants

Lepidodendrales or arborescent lycophytes are an extinct order 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.

<span class="mw-page-title-main">Woody plant</span> Plant that produces wood and has a hard stem

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.

<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, photosynthesis takes place here, stores nutrients, and produces new living tissue. The stem can also be called halm or haulm or culms.

<span class="mw-page-title-main">Cambium</span> Layer of plant tissue with cells for growth

A cambium, in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from which phloem, xylem, or cork grows by division, resulting in secondary thickening. It forms parallel rows of cells, which result in secondary tissues.

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  45. ↑ Paulo Fernandes (3 January 2011). "j.g. pausas' blog " Bark thickness: a world record?". Jgpausas.blogs.uv.es. doi:10.1016/j.foreco.2010.07.010. hdl: 10261/32660 . S2CID   85573811 . Retrieved 2 January 2012.{{cite journal}}: Cite journal requires |journal= (help)
  46. ↑ Ibrahim, Mohammed Auwal; Mohammed, Aminu; Isah, Murtala Bindawa; Aliyu, Abubakar Babando (2014). "Anti-trypanosomal activity of African medicinal plants: A review update". Journal of Ethnopharmacology . 154 (1). International Society of Ethnopharmacology (ISE): 26–54. doi:10.1016/j.jep.2014.04.012. ISSN   0378-8741. PMID   24742753.
  47. ↑ Myking, T.; Hertzberg, A.; Skrøppa, T. (2005). "History, manufacture and properties of lime bast cordage in northern Europe". Forestry. 78 (1): 65–71. doi: 10.1093/forestry/cpi006 .
  48. ↑ Duran-Reynals, Marie Louise de Ayala. 1946. The Fever Bark Tree; the Pageant of Quinine. Garden City, N.Y.: Doubleday.
  49. ↑ Markham, Clements R. 1880. Peruvian Bark. A Popular Account of the Introduction of Chinchona Cultivation into British India. London: J. Murray.
  50. ↑ Harkin, John M. (1971). Bark and Its Possible Uses. Forest Products Laboratory, U.S. Forest Service.
  51. ↑ "Archived copy" (PDF). Archived from the original (PDF) on 11 April 2008. Retrieved 30 January 2008.{{cite web}}: CS1 maint: archived copy as title (link)

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