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. [2] 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 [ s ], derives from its lenticular (lens-like) shape. [3] The shape of lenticels is one of the characteristics used for tree identification. [4]
Before there was much evidence for the existence and functionality of lenticels, the fossil record has shown the first primary mechanism of aeration in early vascular plants to be the stomata. [5]
However, in woody plants, while the respiratory function of stomata is retained in the living epidermis of leaves and green stems, that function is lost where the epidermis of trunks and branches is displaced by vascular and cork cambial activity and by secondary growth. In such parts the entire epidermis may be shed as it is replaced by a suberized periderm or bark in which the respiratory functions of the stomata may be replaced by lenticels, at least until the bark becomes too thick.
The extinct arboreal plants of the genera Lepidodendron and Sigillaria were the first to have distinct aeration structures that rendered these modifications. "Parichnoi" (singular: parichnos) are canal-like structures that, in association with foliar traces of the stem, connected the stem's outer and middle cortex to the mesophyll of the leaf. Parichnoi were thought to eventually give rise to lenticels as they helped solve the issue of long-range oxygen transport in these woody plants during the Carboniferous period. They also acquired secondary connections as they evolved to become transversely elongated to efficiently aerate the maximum number of vertical rays as well as the central core tissue of the stem. [6] The evolutionary significance of parichnoi was their functionality in the absence of cauline stomata, where they can also be affected and destroyed by pressure similar to what can damage to stomatal tissue. Evidently, in both conifers and Lepidodendroids, the parichnoi, as the primary lenticular structure, appear as paired structures on either side of leaf scars. The development and increase in the number of these primitive lenticels were key to providing a system that was open for aeration and gas exchange in these plants. [7]
In plant bodies that produce secondary growth, lenticels promote gas exchange of oxygen, carbon dioxide, and water vapor. [8] Lenticel formation usually begins beneath stomatal complexes during primary growth preceding the development of the first periderm. The formation of lenticels seem to be directly related to the growth and strength of the shoot and on the hydrose of the tissue, which refers to the internal moisture. [9] As stems and roots mature lenticel development continues in the new periderm (for example, periderm that forms at the bottom of cracks in the bark).[ citation needed ]
Lenticels are found as raised circular, oval, or elongated areas on stems and roots. In woody plants, lenticels commonly appear as rough, cork-like structures on young branches. Underneath them, porous tissue creates a number of large intercellular spaces between cells. This tissue fills the lenticel and arises from cell division in the phellogen or substomatal ground tissue. Discoloration of lenticels may also occur, such as in mangoes, that may be due to the amount of lignin in cell walls. [10] [11]
In oxygen deprived conditions, making respiration a daily challenge, different species may possess specialized structures where lenticels can be found. For example, in a common mangrove species, lenticels appear on pneumatophores (specialized roots), where the parenchyma cells that connect to the aerenchyma structure increase in size and go through cell division. [12] In contrast, lenticels in grapes are located on the pedicels and act as a function of temperature. If they are blocked, hypoxia and successive ethanol accumulation may result and lead to cell death. [13]
Lenticels are also present on many fruits, quite noticeably on many apples and pears. On European pears, they can serve as an indicator of when to pick the fruit, as light lenticels on the immature fruit darken and become brown and shallow from the formation of cork cells. [14] [15] Certain bacterial and fungal infections can penetrate fruits through their lenticels, with susceptibility sometimes increasing with its age. [16]
While the term lenticel is usually associated with the breakage of periderm tissue that is associated with gas exchange, it also refers to the lightly colored spots found on apples (a type of pome fruit). "Lenticel" seems to be the most appropriate term to describe both structures mentioned in light of their similar function in gas exchange. Pome lenticels can be derived from no longer functioning stomata, epidermal breaks from the removal of trichomes, and other epidermal breaks that usually occur in the early development of young pome fruits. The closing of pome lenticels can arise when the cuticle over the stomata opening or the substomatal layer seals. Closing can also begin if the substomatal cells become suberized, like cork. The number of lenticels usually varies between the species of apples, where the range may be from 450 to 800 or from 1500 to 2500 in Winesap and Spitzenburg apples, respectively. This wide range may be due to the water availability during the early stages of development of each apple type. [17]
"Lenticel breakdown" is a global skin disorder of apples in which lenticels develop dark 1–8 mm diameter pits shortly after processing and packing. [18] [19] It is most common on the 'Gala' (Malus × domestica) variety, particularly the 'Royal Gala', and also occurs in 'Fuji', 'Granny Smith', 'Golden Delicious', and 'Delicious' varieties. [18] [19] It is more common in arid regions, and is thought to be related to relative humidity and temperature. [18] [19] The effect can be mitigated by spraying the fruit with lipophilic coatings prior to harvest. [18]
Lenticels are also present on potato tubers. [20]
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. 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 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.
In botany, a stoma, also called a stomate, is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange between the internal air spaces of the leaf and the atmosphere. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that regulate the size of the stomatal opening.
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.
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.
In cell biology, 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.
Aerial roots are roots growing above the ground. They are often adventitious, i.e. formed from nonroot tissue. They are found in diverse plant species, including epiphytes such as orchids (Orchidaceae), tropical coastal swamp trees such as mangroves, banyan figs, the warm-temperate rainforest rata, and pohutukawa trees of New Zealand. Vines such as common ivy and poison ivy also have aerial roots.
Sclereids are a reduced form of sclerenchyma cells with highly thickened, lignified cellular walls that form small bundles of durable layers of tissue in most plants. The presence of numerous sclereids form the cores of apples and produce the gritty texture of guavas.
Suberin is a lipophilic, complex polyester biopolymer of plants, composed of long-chain fatty acids called suberin acids and glycerol. Suberin, interconnected with cutins and lignins, also complex macromolecules, form a protective barrier in the epidermal and peridermal cell walls of higher plants. 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.
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.
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.
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.
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.
A plant cuticle is a protecting film covering the outermost skin layer (epidermis) of leaves, young shoots and other aerial plant organs that have no periderm. The film consists of lipid and hydrocarbon polymers infused with wax, and is synthesized exclusively by the epidermal cells.
Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally, it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century, plant anatomy has been considered a separate field referring only to internal plant structure. Plant anatomy is now frequently investigated at the cellular level, and often involves the sectioning of tissues and microscopy.
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.
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, engages in photosynthesis, stores nutrients, and produces new living tissue. The stem can also be called the culm, halm, haulm, stalk, or thyrsus.
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.
Hypericum sechmenii, or Seçmen's St John's wort, is a rare species of flowering plant of the St John's wort family (Hypericaceae) that is found in the Eskişehir Province of central Turkey. It was first described in 2009 by Turkish botanists Atila Ocak and Onur Koyuncu, who named the species in honor of Özcan Seçmen, a fellow botanist. They assigned the species to the genus Hypericum, and Norman Robson later placed H. sechmenii into the section Adenosepalum.