In plant morphology, thorns, spines, and prickles, and in general spinose structures (sometimes called spinose teeth or spinose apical processes), are hard, rigid extensions or modifications of leaves, roots, stems, or buds with sharp, stiff ends, and generally serve the same function: physically defending plants against herbivory.
In common language, the terms are used more or less interchangeably, but in botanical terms, thorns are derived from shoots (so that they may or may not be branched, they may or may not have leaves, and they may or may not arise from a bud), [1] [2] [3] [4] spines are derived from leaves (either the entire leaf or some part of the leaf that has vascular bundles inside, like the petiole or a stipule), [1] [2] [3] [4] and prickles are derived from epidermis tissue (so that they can be found anywhere on the plant and do not have vascular bundles inside [4] ). [1] [2] [3]
Leaf margins may also have teeth, and if those teeth are sharp, they are called spinose teeth on a spinose leaf margin [1] [2] (some authors consider them a kind of spine [2] ). On a leaf apex, if there is an apical process (generally an extension of the midvein), and if it is especially sharp, stiff, and spine-like, it may be referred to as spinose or as a pungent apical process [1] (again, some authors call them a kind of spine [2] ). When the leaf epidermis is covered with very long, stiff trichomes (more correctly called bristles in this case; [1] for some authors a kind of prickle [2] ), it may be referred to as a hispid vestiture; [1] [2] [3] if the trichomes are stinging trichomes, it may be called a urent vestiture. [1]
There can be found also spines or spinose structures derived from roots. [5]
The predominant function of thorns, spines, and prickles is deterring herbivory in a mechanical form. For this reason, they are classified as physical or mechanical defenses, as opposed to chemical defenses.
Not all functions of spines or glochids are limited to defense from physical attacks by herbivores and other animals. In some cases, spines have been shown to shade or insulate the plants that grow them, thereby protecting them from extreme temperatures. For example, saguaro cactus spines shade the apical meristem in summer, and in members of the Opuntioideae, glochids insulate the apical meristem in winter.
Agrawal et al. (2000) found that spines seem to have little effect on specialist pollinators, on which many plants rely in order to reproduce. [6]
Pointing or spinose processes can broadly be divided by the presence of vascular tissue: thorns and spines are derived from shoots and leaves respectively, and have vascular bundles inside, whereas prickles (like rose prickles) do not have vascular bundles inside, so that they can be removed more easily and cleanly than thorns and spines.
Thorns are modified branches or stems. They may be simple or branched.
Spines are modified leaves, stipules, or parts of leaves, such as extensions of leaf veins. Some authors prefer not to distinguish spines from thorns because, like thorns, and unlike prickles, they commonly contain vascular tissue. [7]
Spines are variously described as petiolar spines (as in Fouquieria ), leaflet spines (as in Phoenix ), or stipular spines (as in Euphorbia ), all of which are examples of spines developing from a part of a leaf containing the petiole, midrib, or a secondary vein. [1] The plants of the cactus family are particularly well known for their dense covering of spines. Some cacti have also glochids (or glochidia, singular glochidium) – a particular kind of spine of different origin, which are smaller and deciduous with numerous retrose barbs along its length (as found in areoles of Opuntia ). [1]
Prickles are comparable to hairs but can be quite coarse (for example, rose prickles). They are extensions of the cortex and epidermis. [8] [9] Technically speaking, many plants commonly thought of as having thorns or spines actually have prickles. Roses, for instance, have prickles. [7] While the position of thorns and spines are known positively to be controlled by phyllotaxis, the positioning of prickles appears to be truly random. If not, then by a phyllotaxis so arcane as to give the appearance of randomness.[ citation needed ]
A study published for peer review to the journal Science concluded that plants with these types of prickles have been identified as sharing a common gene family. [10]
Other similar structures are spinose teeth, spinose apical processes, and trichomes. Trichomes, in particular, are distinct from thorns, spines, and prickles in that they are much smaller (often microscopic) outgrowths of epidermal tissue, and they are less rigid and more hair-like in appearance; they typically consist of just a few cells of the outermost layer of epidermis, whereas prickles may include cortex tissue. Trichomes are often effective defenses against small insect herbivores; thorns, spines, and prickles are usually only effective against larger herbivores like birds and mammals.
Spinescent is a term describing plants that bear any sharp structures that deter herbivory. It also can refer to the state of tending to be or become spiny in some sense or degree, as in: "... the division of the African acacias on the basis of spinescent stipules versus non-spinescent stipules..." [11]
There are also spines derived from roots, like the ones on the trunk of the "Root Spine Palms" ( Cryosophila spp.). The trunk roots of Cryosophila guagara grow downwards to a length of 6–12 cm, then stop growing and transform into a spine. [5] The anatomy of crown roots on this species (roots among the bases of the living fronds) also alters during their life. [5] They initially grow upwards and then turn down and finally they, too, become spinous. [5] Lateral roots on these two types of roots, as well as those on the stilt roots on this species, also become spinous. [5] Some authors believe that some of these short spiny laterals have a ventilating function so they are 'pneumorhizae'. [5] Short spiny laterals that may have a ventilating function may also be found on roots of Iriartea exorrhiza . [5]
There are also spines that function as pneumorhizae on the palm Euterpe oleracea . [5] In Cryosophila nana (formerly Acanthorhiza aculeata), there are spiny roots; some authors prefer to term these "root spines" if the length of the root is less than 10x the thickness and "spine roots" if the length is more than 10x the thickness. [5] Adventitious spiny roots have also been described on the trunks of dicotyledonous trees from tropical Africa (e.g. Euphorbiaceae, as in Macaranga barteri , Bridelia micrantha and B. pubescens ; Ixonanthaceae, Sterculiaceae), and may also be found protecting perennating organs such as tubers and corms (e.g. Dioscorea prehensilis -Dioscoreaceae- and Moraea spp. -Iridaceae- respectively). [5] Short root spines cover the tuberous base of the epiphytic ant-plant Myrmecodia tuberosa (Rubiaceae), these probably give protection to ants which inhabit chambers within the tuber as they wander over the plant's surface. (Jackson 1986 [5] and references therein). In many respects, the pattern of spine formation is similar to that which occurs in the development of thorns from lateral shoots. (Jackson 1986 [5] and references therein).
It has been proposed that thorny structures may have first evolved as a defense mechanism in plants growing in sandy environments that provided inadequate resources for fast regeneration of damage. [12] [13]
Spinose structures occur in a wide variety of ecologies, and their morphology also varies greatly. They occur as:
Some thorns are hollow and act as myrmecodomatia; others (e.g. in Crataegus monogyna ) bear leaves. The thorns of many species are branched (e.g. in Crataegus crus-galli and Carissa macrocarpa ).
Plants bearing thorns, spines, or prickles are often used as a defense against burglary, being strategically planted below windows or around the entire perimeter of a property. [17] They also have been used to protect crops and livestock against marauding animals. Examples include hawthorn hedges in Europe, agaves or ocotillos in the Americas and in other countries where they have been introduced, Osage orange in the prairie states of the US, and Sansevieria in Africa. [18] [ page needed ]
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthparts adapted to rasping or grinding. Horses and other herbivores have wide flat teeth that are adapted to grinding grass, tree bark, and other tough plant material.
Trichomes are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant is an indumentum, and the surface bearing them is said to be pubescent.
In botany, a stipule is an outgrowth typically borne on both sides of the base of a leafstalk. Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many species they may be inconspicuous —or sometimes entirely absent, and the leaf is then termed exstipulate. The word stipule was coined by Linnaeus from Latin stipula, straw, stalk.
The following terms are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple or compound. The edge of the leaf may be regular or irregular, may be smooth or bearing hair, bristles or spines. For more terms describing other aspects of leaves besides their overall morphology see the leaf article.
Kalanchoe beharensis is a plant species in the succulent genus Kalanchoe, and the family Crassulaceae. Kalanchoe beharensis is native to Madagascar known by local names mongy, rongy and tavitavy.
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.
Vachellia cornigera, commonly known as bullhorn acacia, is a swollen-thorn tree and Myrmecophyte native to Mexico and Central America. The common name of "bullhorn" refers to the enlarged, hollowed-out, swollen thorns that occur in pairs at the base of leaves, and resemble the horns of a steer. In Yucatán it is called "subín", in Panamá the locals call them "cachito". The trees are commonly found in wet lowlands
Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide herbivore protection. Common nectar-consuming pollinators include mosquitoes, hoverflies, wasps, bees, butterflies and moths, hummingbirds, honeyeaters and bats. Nectar plays a crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior is largely responsible for the divergent evolution of the African honey bee, A. m. scutellata and the western honey bee.
Plant defense against herbivory or host-plant resistance (HPR) is a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Many plants produce secondary metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores or reduce plant digestibility. Another defensive strategy of plants is changing their attractiveness. Plants can sense being touched, and they can respond with strategies to defend against herbivores. To prevent overconsumption by large herbivores, plants alter their appearance by changing their size or quality, reducing the rate at which they are consumed.
Vachellia collinsii, previously Acacia collinsii, is a species of flowering plant native to Central America and parts of Africa.
Herbivores are dependent on plants for food, and have coevolved mechanisms to obtain this food despite the evolution of a diverse arsenal of plant defenses against herbivory. Herbivore adaptations to plant defense have been likened to "offensive traits" and consist of those traits that allow for increased feeding and use of a host. Plants, on the other hand, protect their resources for use in growth and reproduction, by limiting the ability of herbivores to eat them. Relationships between herbivores and their host plants often results in reciprocal evolutionary change. When a herbivore eats a plant it selects for plants that can mount a defensive response, whether the response is incorporated biochemically or physically, or induced as a counterattack. In cases where this relationship demonstrates "specificity", and "reciprocity", the species are thought to have coevolved. The escape and radiation mechanisms for coevolution, presents the idea that adaptations in herbivores and their host plants, has been the driving force behind speciation. The coevolution that occurs between plants and herbivores that ultimately results in the speciation of both can be further explained by the Red Queen hypothesis. This hypothesis states that competitive success and failure evolve back and forth through organizational learning. The act of an organism facing competition with another organism ultimately leads to an increase in the organism's performance due to selection. This increase in competitive success then forces the competing organism to increase its performance through selection as well, thus creating an "arms race" between the two species. Herbivores evolve due to plant defenses because plants must increase their competitive performance first due to herbivore competitive success.
A bur is a seed or dry fruit or infructescence that has hooks or teeth. The main function of the bur is to spread the seeds of the bur plant, often through epizoochory. The hooks of the bur are used to latch onto fur or fabric, enabling the bur – which contain seeds – to be transported to another location for dispersal. Another use for the spines and hooks are physical protection against herbivores. Their ability to stick to animals and fabrics has shaped their reputation as bothersome.
This page provides a glossary of plant morphology. Botanists and other biologists who study plant morphology use a number of different terms to classify and identify plant organs and parts that can be observed using no more than a handheld magnifying lens. This page provides help in understanding the numerous other pages describing plants by their various taxa. The accompanying page—Plant morphology—provides an overview of the science of the external form of plants. There is also an alphabetical list: Glossary of botanical terms. In contrast, this page deals with botanical terms in a systematic manner, with some illustrations, and organized by plant anatomy and function in plant physiology.
Vachellia drepanolobium, more commonly known as Acacia drepanolobium or whistling thorn, is a swollen-thorn acacia native to East Africa. The whistling thorn grows up to 6 meters tall. It produces a pair of straight spines at each node, some of which have large bulbous bases. These swollen spines are naturally hollow and occupied by any one of several symbiotic ant species. The common name of the plant is derived from the observation that when wind blows over bulbous spines in which ants have made entry and exit holes, they produce a whistling noise.
This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.
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, or thyrsus.
A leaf is a principal appendage of the stem of a vascular plant, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, flower, and fruit collectively form the shoot system. In most leaves, the primary photosynthetic tissue is the palisade mesophyll and is located on the upper side of the blade or lamina of the leaf but in some species, including the mature foliage of Eucalyptus, palisade mesophyll is present on both sides and the leaves are said to be isobilateral. Most leaves are flattened and have distinct upper (adaxial) and lower (abaxial) surfaces that differ in color, hairiness, the number of stomata, the amount and structure of epicuticular wax and other features. Leaves are mostly green in color due to the presence of a compound called chlorophyll which is essential for photosynthesis as it absorbs light energy from the sun. A leaf with lighter-colored or white patches or edges is called a variegated leaf.
Plants and herbivores have co-evolved together for 350 million years. Plants have evolved many defense mechanisms against insect herbivory. Such defenses can be broadly classified into two categories: (1) permanent, constitutive defenses, and (2) temporary, inducible defenses. Both types are achieved through similar means but differ in that constitutive defenses are present before an herbivore attacks, while induced defenses are activated only when attacks occur. In addition to constitutive defenses, initiation of specific defense responses to herbivory is an important strategy for plant persistence and survival.
In botany, a fascicle is a bundle of leaves or flowers growing crowded together; alternatively the term might refer to the vascular tissues that supply such an organ with nutrients. However, vascular tissues may occur in fascicles even when the organs they supply are not fascicled.
Plants are constantly exposed to different stresses that result in wounding. Plants have adapted to defend themselves against wounding events, like herbivore attacks or environmental stresses. There are many defense mechanisms that plants rely on to help fight off pathogens and subsequent infections. Wounding responses can be local, like the deposition of callose, and others are systemic, which involve a variety of hormones like jasmonic acid and abscisic acid.
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