Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is 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 is the liberation of a fungal

spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...

. 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 and ...

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

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 in American English and Canadian English, 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 ( Sou ...

, 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 new individual organism or offspring and initiate its development. Proce ...

, 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. Reproduction is a fundamental feature of all known life; each individual or ...

. Most deciduous plants drop their leaves by abscission before winter, whereas 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 Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...

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 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, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...

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.Williams, A.G., & T.G. Whitham (1986). Premature leaf abscission: an induced plant defense against gall aphids. Ecology, 67(6), 1619-1627.

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 (also chlorophyl) 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 allow plants to a ...

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 hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

, nucleic acids,

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, respo ...

, and certain

plant hormones Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pat ...

.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 to change colors. Carotenoids 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 Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...

cells, which produce and inject suberin 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 rigidity ...

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 Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...

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 Cytokinesis () is the part of the cell division process during ...

, 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 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

In deciduous trees, an abscission zone, also called a separation zone, is formed at the base of the petiole. 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 aphids, 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 (ROS) are generated by plants during times of stress (

biotic Biotics describe living or once living components of a community; for example organisms, such as animals and plants. Biotic may refer to: *Life, the condition of living organisms *Biology, the study of life * Biotic material, which is derived from ...

and abiotic) including

UV light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30 PHz) to 400 nm (750 THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...

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

salinity Salinity () is the saltiness or amount of 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 dimensionless and equal ...

. The presence and continuous production of these ROS causes disruption in the homeostasis of the cellular components, leading to

metabolic Metabolism (, from el, μεταβολή ''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 cell ...

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) 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 environmental s ...

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 essenti ...

, 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 double bonds). Ethylene i ...

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. The term ''synergy'' comes from the Attic Greek word συνεργία ' from ', , meaning "working together". History In Christi ...

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 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.

In animals

See:

Moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...

, Autotomy, Claw (Mammals)

References

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External links

Absission layer microscopy
Plant anatomy Plant morphology Plant physiology