Abscission
Encyclopedia
Abscission is a term used in several areas of biology. In plant sciences it most commonly refers to the process by which a plant
drops one or more of its parts, such as a leaf
, fruit
, flower
or seed
. In zoology
the term may be used to describe the intentional shedding of a body part, such as the shedding of a claw
husk, or the autotomy
of a tail to evade a predator. In mycology
it may refer to the liberation of a fungal spore
. In cell biology abscission refers to the separation of two daughter cells at the completion of cytokinesis
.
, or for the purposes of reproduction. Most deciduous
plants drop their leaves by abscission before winter, while evergreen
plants continuously abscise their leaves. Another form of abscission is fruit drop, when a plant abscises fruit while still immature, in order to conserve resources needed to bring the remaining fruit to maturity. If a leaf is damaged a plant may also abscise it to conserve water
or photosynthetic efficiency, depending on the 'costs' to the plant as a whole. The abscission layer is a greenish grayish color.
Abscission can also occur in premature leaves as a means of plant defense. Premature leaf abscission has been shown to occur in response to infestation by gall aphids. By abscising leaves that have been made host to aphid galls, plants have been shown to massively diminish the pest population, as 98% of aphids in abscised galls died. The abscission is selective, and the chance of dropping leaves increases as the number of galls increase. A leaf with three or more galls was four times more likely to abscise than a leaf with one, and 20 times as likely to be dropped as a leaf without any galls.
trees, an abscission zone, also called a separation zone, is formed at the base of the petiole. It is composed of a top layer which has cells with weak walls, and a bottom layer which expands in the autumn, breaking the weak walls of the cells in the top layer. This allows the leaf to be shed.
In woody plant
s, an abscission layer is formed composed of parenchyma cells bounded on both sides with cork. This layer is found at the base of the leaf petioles in woody angiosperms and gymnosperms and because of the disintegration of the parenchyma layer, the organ, such as a leaf or bark, is separated from the parent plant. Abscission is a natural process of plant growth induced by the plant, in contrast to decaying
or falling off due to other causes.
production in leaves due to decreased sunlight in the fall explains why some leaves turn yellow. However, the yellow color can attract aphid
s, so some trees turn the leaves red instead by injecting a bright pigment. The loss of chlorophyll may also contribute to the abscission process.
, singlet oxygen
and radical hydroxyl) are generated by plants during times of stress (biotic
and abiotic) including UV light, cool temperatures, excessive light, pathogens, parasites, and high salinity
. The presence and continuous production of these ROS causes disruption in the homeostasis
of the cellular components, leading to metabolic dysfunction and expression of cell wall degrading enzymes (WDE’s).
(Indol-3-acetic acid or IAA, a plant hormone) and ethylene
have been implicated as prominent regulators of abscission signaling. The two compounds work in a synergistic fashion. As the IAA levels decrease, the flux of IAA to the abscission zone is reduced. Exhaustion of IAA makes the abscission zone sensitive to ethylene. When the plant is then exposed to ethylene, gene expression of cell wall degrading enzymes such as cellulase and polygalacturonase
are activated. However, this is not to say that ethylene directly activates WDE gene expression, because the elements responsible for detecting ethylene have not been found in the gene’s promoter region.
While researchers originally believed abscisic acid
to be the hormone that stimulated abscission (for which the hormone was named), it was later proven that it does not play a primary role.
Dwindling auxin levels have also been implicated in autumn-leaf color change.
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
drops one or more of its parts, such as a leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
, fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
, flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
or seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
. In zoology
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...
the term may be used to describe the intentional shedding of a body part, such as the shedding of a claw
Claw
A claw is a curved, pointed appendage, found at the end of a toe or finger in most mammals, birds, and some reptiles. However, the word "claw" is also often used in reference to an invertebrate. Somewhat similar fine hooked structures are found in arthropods such as beetles and spiders, at the end...
husk, or the autotomy
Autotomy
Autotomy or self amputation is the act whereby an animal severs one or more of its own appendages, usually as a self-defense mechanism designed to elude a predator's grasp...
of a tail to evade a predator. In mycology
Mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans as a source for tinder, medicinals , food and entheogens, as well as their dangers, such as poisoning or...
it may refer to the liberation of a fungal spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
. In cell biology abscission refers to the separation of two daughter cells at the completion of cytokinesis
Cytokinesis
Cytokinesis is the process in which the cytoplasm of a single eukaryotic cell is divided to form two daughter cells. It usually initiates during the late stages of mitosis, and sometimes meiosis, splitting a binucleate cell in two, to ensure that chromosome number is maintained from one generation...
.
Function
A plant will abscise a part either to discard a member that is no longer necessary, such as a leaf during autumn, or a flower following fertilisationFertilisation
Fertilisation is the fusion of gametes to produce a new organism. In animals, the process involves the fusion of an ovum with a sperm, which eventually leads to the development of an embryo...
, or for the purposes of reproduction. Most deciduous
Deciduous
Deciduous means "falling off at maturity" or "tending to fall off", and is typically used in reference to trees or shrubs that lose their leaves seasonally, and to the shedding of other plant structures such as petals after flowering or fruit when ripe...
plants drop their leaves by abscission before winter, while evergreen
Evergreen
In botany, an evergreen plant is a plant that has leaves in all seasons. This contrasts with deciduous plants, which completely lose their foliage during the winter or dry season.There are many different kinds of evergreen plants, both trees and shrubs...
plants continuously abscise their leaves. Another form of abscission is fruit drop, when a plant abscises fruit while still immature, in order to conserve resources needed to bring the remaining fruit to maturity. If a leaf is damaged a plant may also abscise it to conserve water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...
or photosynthetic efficiency, depending on the 'costs' to the plant as a whole. The abscission layer is a greenish grayish color.
Abscission can also occur in premature leaves as a means of plant defense. Premature leaf abscission has been shown to occur in response to infestation by gall aphids. By abscising leaves that have been made host to aphid galls, plants have been shown to massively diminish the pest population, as 98% of aphids in abscised galls died. The abscission is selective, and the chance of dropping leaves increases as the number of galls increase. A leaf with three or more galls was four times more likely to abscise than a leaf with one, and 20 times as likely to be dropped as a leaf without any galls.
Structural mechanism
In deciduousDeciduous
Deciduous means "falling off at maturity" or "tending to fall off", and is typically used in reference to trees or shrubs that lose their leaves seasonally, and to the shedding of other plant structures such as petals after flowering or fruit when ripe...
trees, an abscission zone, also called a separation zone, is formed at the base of the petiole. It is composed of a top layer which has cells with weak walls, and a bottom layer which expands in the autumn, breaking the weak walls of the cells in the top layer. This allows the leaf to be shed.
In woody plant
Woody plant
A woody plant is a plant that uses wood as its structural tissue. These are typically perennial plants whose stems and larger roots are reinforced with wood produced adjacent to the vascular tissues. The main stem, larger branches, and roots of these plants are usually covered by a layer of...
s, an abscission layer is formed composed of parenchyma cells bounded on both sides with cork. This layer is found at the base of the leaf petioles in woody angiosperms and gymnosperms and because of the disintegration of the parenchyma layer, the organ, such as a leaf or bark, is separated from the parent plant. Abscission is a natural process of plant growth induced by the plant, in contrast to decaying
Decomposition
Decomposition is the process by which organic material is broken down into simpler forms of matter. The process is essential for recycling the finite matter that occupies physical space in the biome. Bodies of living organisms begin to decompose shortly after death...
or falling off due to other causes.
Lack of chlorophyll as a trigger
The reduction of chlorophyllChlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...
production in leaves due to decreased sunlight in the fall explains why some leaves turn yellow. However, the yellow color can attract aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
s, so some trees turn the leaves red instead by injecting a bright pigment. The loss of chlorophyll may also contribute to the abscission process.
Chemical mechanism
A variety of reactive oxygen species or "ROS" (namely H2O2, but also superoxideSuperoxide
A superoxide, also known by the obsolete name hyperoxide, is a compound that possesses the superoxide anion with the chemical formula O2−. The systematic name of the anion is dioxide. It is important as the product of the one-electron reduction of dioxygen O2, which occurs widely in nature...
, singlet oxygen
Singlet oxygen
Singlet oxygen is the common name used for the diamagnetic form of molecular oxygen , which is less stable than the normal triplet oxygen. Because of its unusual properties, singlet oxygen can persist for over an hour at room temperature, depending on the environment...
and radical hydroxyl) are generated by plants during times of stress (biotic
Biotic component
Biotic components are the living things that shape an ecosystem. A biotic factor is any living component that affects another organism, including animals that consume the organism in question, and the living food that the organism consumes. Each biotic factor needs energy to do work and food for...
and abiotic) including UV light, cool temperatures, excessive light, pathogens, parasites, and high salinity
Salinity
Salinity is the saltiness or dissolved salt content of a body of water. It is a general term used to describe the levels of different salts such as sodium chloride, magnesium and calcium sulfates, and bicarbonates...
. The presence and continuous production of these ROS causes disruption in the homeostasis
Homeostasis
Homeostasis is the property of a system that regulates its internal environment and tends to maintain a stable, constant condition of properties like temperature or pH...
of the cellular components, leading to metabolic dysfunction and expression of cell wall degrading enzymes (WDE’s).
Hormone involvement
AuxinAuxin
Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins have a cardinal role in coordination of many growth and behavioral processes in the plant's life cycle and are essential for plant body development. Auxins and their role in plant growth were first described by...
(Indol-3-acetic acid or IAA, a plant hormone) and ethylene
Ethylene
Ethylene is a gaseous organic compound with the formula . It is the simplest alkene . Because it contains a carbon-carbon double bond, ethylene is classified as an unsaturated hydrocarbon. Ethylene is widely used in industry and is also a plant hormone...
have been implicated as prominent regulators of abscission signaling. The two compounds work in a synergistic fashion. As the IAA levels decrease, the flux of IAA to the abscission zone is reduced. Exhaustion of IAA makes the abscission zone sensitive to ethylene. When the plant is then exposed to ethylene, gene expression of cell wall degrading enzymes such as cellulase and polygalacturonase
Polygalacturonase
Polygalacturonase is an enzyme produced in plants which is involved in the ripening process, and by some bacteria and fungi which is involved in the rotting process. PGs degrades polygalacturonan present in the cell walls of plants by hydrolysis of the glycosidic bonds that link galacturonic acid...
are activated. However, this is not to say that ethylene directly activates WDE gene expression, because the elements responsible for detecting ethylene have not been found in the gene’s promoter region.
While researchers originally believed abscisic acid
Abscisic acid
Abscisic acid , also known as abscisin II and dormin, is a plant hormone. ABA functions in many plant developmental processes, including bud dormancy. It is degraded by the enzyme -abscisic acid 8'-hydroxylase.-Function:...
to be the hormone that stimulated abscission (for which the hormone was named), it was later proven that it does not play a primary role.
Dwindling auxin levels have also been implicated in autumn-leaf color change.