Abscission () is the shedding of various parts of an

organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...

, such as a

plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...

dropping a

leaf A leaf (: leaves) is a principal appendage of the plant stem, stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leav ...

fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...

flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...

, or

seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...

. In

zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...

, abscission is the intentional shedding of a body part, such as the shedding of a

claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or Arthro ...

, husk, or the

autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or 'self-amputation', is the behaviour whereby an animal sheds or discards an appendage, usually as a self-defense mechanism to elude a predator's grasp ...

of a tail to evade a predator. In

mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their Taxonomy (biology), taxonomy, genetics, biochemistry, biochemical properties, and ethnomycology, use by humans. Fungi can be a source of tinder, Edible ...

, it is the liberation of a fungal

spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...

. In

cell biology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living an ...

, abscission refers to the separation of two daughter cells at the completion of

cytokinesis Cytokinesis () is the part of the cell division process and part of mitosis during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division ...

In plants

Function

A plant will abscise a part either to discard a member that is no longer necessary, such as a leaf during

autumn Autumn, also known as fall (especially in US & Canada), is one of the four temperate seasons on Earth. Outside the tropics, autumn marks the transition from summer to winter, in September (Northern Hemisphere) or March ( Southern Hemisphe ...

, or a flower following

fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or of ...

, or for the purposes of

reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...

. Most

deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...

plants drop their leaves by abscission before

winter Winter is the coldest and darkest season of the year in temperate and polar climates. It occurs after autumn and before spring. The tilt of Earth's axis causes seasons; winter occurs when a hemisphere is oriented away from the Sun. Dif ...

, whereas

evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...

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 is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

efficiency, depending on the 'costs' to the plant as a whole. The abscission layer is a greenish-greyish 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 Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...

. 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 Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or wart ...

died. The abscission is selective, and the chance of dropping leaves increases as the number of galls increases. 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.

Process

Abscission occurs in a series of three events: 1) resorption, 2) protective layer formation, and 3) detachment.Addicott, F.T. 1982. Abscission. University of California Press, London, England. Steps 2 and 3 may occur in either order depending on the species. Pomegranate Bonsai in Fall

Resorption

Resorption involves degrading

chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...

to extract the majority of its nutrients.Keskitalo, J., G. Bergguist, P. Gardestrom, and S. Jansson. 2005. A Cellular Timetable of Autumn Senescence. Plant Phys. 139 : 1635-1648. Nitrogen is found in chlorophyll and is often a

limiting nutrient A limiting factor is a variable of a system that causes a noticeable change in output or another measure of a type of system. The limiting factor is in a pyramid shape of organisms going up from the producers to consumers and so on. A factor not l ...

for plants, which need large quantities to form

amino acids Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the Proteinogenic amino acid, 22 α-amino acids incorporated into p ...

nucleic acids Nucleic acids are large biomolecules that are crucial in all cells and viruses. They are composed of nucleotides, which are the monomer components: a 5-carbon sugar, a phosphate group and a nitrogenous base. The two main classes of nucleic a ...

proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, re ...

, and certain

plant hormones Plant hormones (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of Organ (anat ...

.Hopkins, W.G. and N.P.A. Huner. 2009. Introduction to Plant Physiology. Fourth edition. Wiley & Sons, Hoboken, NJ. Once nitrogen and other nutrients have been extracted from chlorophyll, the nutrients will travel to other tissues of the plant. Resorption is what causes leaves in the

fall Autumn, also known as fall (especially in US & Canada), is one of the four temperate seasons on Earth. Outside the tropics, autumn marks the transition from summer to winter, in September (Northern Hemisphere) or March ( Southern Hemispher ...

to change colors.

Carotenoids Carotenoids () are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and Fungus, fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips ...

in the leaves are slower to degrade than chlorophyll, so autumn leaves appear yellow and orange.

Protective layer formation

Cells under the abscission zone divide and form a layer of cork cells.Kozlowski, T.T. 1973. Shedding of Plant Parts. Academic Press, New York, NY. Situated on both sides of the abscission zone are layers of

parenchyma upright=1.6, Lung parenchyma showing damage due to large subpleural bullae. Parenchyma () is the bulk of functional substance in an animal organ such as the brain or lungs, or a structure such as a tumour. In zoology, it is the tissue that ...

cells, which produce and inject

suberin Suberin is a lipophilic, complex polyester biopolymer found in plants. It is composed of long-chain fatty acids (called suberin acids) and glycerol. Suberin is interconnected with cutin and lignin and forms a protective barrier in the epidermal ...

and

lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidit ...

under the abscission zone into the new layer of cork cells. Suberin and lignin create a durable and waterproof layer for the plant once the organ is detached.

Detachment

This step can occur in a variety of ways depending on the species but always occurs at the abscission zone.Solomon, E.P., L.R. Berg., and D.W. Martin. 2011. Biology. Ninth edition, Brooks/Cole, Belmont, CA. Detachment can occur when layers of parenchyma cells secrete cell wall

enzymes An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as pro ...

to self-digest the

middle lamella The middle lamella is a layer that cements together the primary cell walls of two adjoining plant cells. It is the first formed layer to be deposited at the time of cytokinesis. The cell plate that is formed during cell division itself develops ...

, which holds the cell walls together at the abscission zone. This causes the cells of the abscission zone to break apart and the leaf or other plant part to fall off. Another way detachment occurs is through

imbibition Imbibition is a special type of diffusion that takes place when liquid is absorbed by solids-Colloid, colloids causing an increase in volume. Water surface potential movement takes place along a concentration gradient; some dry materials absorb ...

of water. The plant cells at the abscission zone will take in a large amount of water, swell, and eventually burst, making the organ fall off. Once detached, the protective layer of cork will be exposed.

Mechanisms

Structural

trees, an abscission zone, also called a separation zone, is formed at the base of the

petiole Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, and ...

. It is composed of a top layer that has cells with weak walls, and a bottom layer that expands in the autumn, breaking the weak walls of the cells in the top layer. This allows the leaf to be shed.

Lack of chlorophyll as a trigger

The reduction of

production in leaves due to decreased sunlight in the autumn explains why some leaves turn yellow. However, the yellow color can attract

aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...

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

A variety of

reactive oxygen species In chemistry and biology, reactive oxygen species (ROS) are highly Reactivity (chemistry), reactive chemicals formed from diatomic oxygen (), water, and hydrogen peroxide. Some prominent ROS are hydroperoxide (H2O2), superoxide (O2−), hydroxyl ...

(ROS) are generated by plants during times of stress ( biotic and

abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them und ...

), including

UV light Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of t ...

, cool temperatures, excessive light, pathogens, parasites, and high

salinity Salinity () is the saltiness or amount of salt (chemistry), salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensio ...

. The presence and continuous production of these ROS causes disruption in the

homeostasis In biology, homeostasis (British English, British also homoeostasis; ) is the state of steady internal physics, physical and chemistry, chemical conditions maintained by organism, living systems. This is the condition of optimal functioning fo ...

of the cellular components, leading to

metabolic Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the ...

dysfunction and expression of cell wall-degrading enzymes (WDEs).Sakamoto, M., I. Munemura, R. Tomita, & K. Kobayashi (2008). Reactive oxygen species in leaf abscission signaling. Plant Signal Behavior, 3(11), 1014-1015.

Hormonal

While researchers originally believed

abscisic acid Abscisic acid (ABA or abscisin II) is a plant hormone. ABA functions in many plant developmental processes, including seed and bud dormancy, the control of organ size and stomatal closure. It is especially important for plants in the response to ...

to be the hormone that stimulates abscission (for which the hormone was named), it was later proven that it does not play a primary role. In fact,

auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essent ...

, a plant hormone, and

ethylene Ethylene (IUPAC name: ethene) is a hydrocarbon which has the formula or . It is a colourless, flammable gas with a faint "sweet and musky" odour when pure. It is the simplest alkene (a hydrocarbon with carbon–carbon bond, carbon–carbon doub ...

have been implicated as prominent regulators of abscission signaling. The two compounds work in a

synergistic Synergy is an interaction or cooperation giving rise to a whole that is greater than the simple sum of its parts (i.e., a non-linear addition of force, energy, or effect). The term ''synergy'' comes from the Attic Greek word συνεργία ' f ...

fashion: As the auxin levels decrease, the flux of auxin to the abscission zone is reduced. Exhaustion of auxin 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 Endo-polygalacturonase (, pectin depolymerase, pectolase, pectin hydrolase, and poly-α-1,4-galacturonide glycanohydrolase; systematic name (1→4)-α-D-galacturonan glycanohydrolase (endo-cleaving)) is an enzyme that hydrolyzes the α-1,4 glycos ...

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. Dwindling auxin levels have also been implicated in autumn-leaf color change.