Plant senescence

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Plant senescence is the process of aging in plants. Plants have both stress-induced and age-related developmental aging. [1] Chlorophyll degradation during leaf senescence reveals the carotenoids, such as anthocyanin and xanthophylls, which are the cause of autumn leaf color in deciduous trees. Leaf senescence has the important function of recycling nutrients, mostly nitrogen, to growing and storage organs of the plant. Unlike animals, plants continually form new organs and older organs undergo a highly regulated senescence program to maximize nutrient export.

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The autumn senescence of Oregon grape leaves is an example of programmed plant senescence. Oregongrapeleaves.jpg
The autumn senescence of Oregon grape leaves is an example of programmed plant senescence.

Hormonal regulation of senescence

Programmed senescence seems to be heavily influenced by plant hormones. The hormones abscisic acid, ethylene, jasmonic acid and salicylic acid are accepted by most scientists as promoters of senescence, but at least one source lists gibberellins, brassinosteroids and strigolactone as also being involved. [2] Cytokinins help to maintain the plant cell and expression of cytokinin biosynthesis genes late in development prevents leaf senescence. [3] A withdrawal of or inability of the cell to perceive cytokinin may cause it to undergo apoptosis or senescence. [4] In addition, mutants that cannot perceive ethylene show delayed senescence. Genome-wide comparison of mRNAs expressed during dark-induced senescence versus those expressed during age-related developmental senescence demonstrate that jasmonic acid and ethylene are more important for dark-induced (stress-related) senescence while salicylic acid is more important for developmental senescence. [5]

Annual versus perennial benefits

Some plants have evolved into annuals which die off at the end of each season and leave seeds for the next, whereas closely related plants in the same family have evolved to live as perennials. This may be a programmed "strategy"[ clarification needed ] for the plants.

The benefit of an annual strategy may be genetic diversity, as one set of genes does continue year after year, but a new mix is produced each year. Secondly, being annual may allow the plants a better survival strategy, since the plant can put most of its accumulated energy and resources into seed production rather than saving some for the plant to overwinter, which would limit seed production.[ citation needed ]

Conversely, the perennial strategy may sometimes be the more effective survival strategy, because the plant has a head start every spring with growing points, roots, and stored energy that have survived through the winter. In trees for example, the structure can be built on year after year so that the tree and root structure can become larger, stronger, and capable of producing more fruit and seed than the year before, out-competing other plants for light, water, nutrients, and space. This strategy will fail when environmental conditions change rapidly. If a certain bug quickly takes advantage and kills all of the nearly identical perennials, then there will be a far lesser chance that a random mutation will slow the bug compared to more diverse annuals.[ citation needed ]

Plant self-pruning

There is a speculative hypothesis on how and why a plant induces part of itself to die off. [2] The theory holds that leaves and roots are routinely pruned off during the growing season whether they are annual or perennial. This is done mainly to mature leaves and roots and is for one of two reasons; either both the leaves and roots that are pruned are no longer efficient enough nutrient acquisition-wise or that energy and resources are needed in another part of the plant because that part of the plant is faltering in its resource acquisition.

This is an oversimplification, in that it is arguable that some shoot and root cells serve other functions than to acquire nutrients. In these cases, whether they are pruned or not would be "calculated" by the plant using some other criteria. It is also arguable that, for example, mature nutrient-acquiring shoot cells would have to acquire more than enough shoot nutrients to support both it and its share of both shoot and root cells that do not acquire sugar and gases whether they are of a structural, reproductive, immature, or just plain, root nature.

The idea that a plant does not impose efficiency demands on immature cells is that most immature cells are part of so-called dormant buds in plants. These are kept small and non-dividing until the plant needs them. They are found in buds, for instance in the base of every lateral stem.

Theory of hormonal induction of senescence

There is little theory on how plants induce themselves to senesce, although it is reasonably widely accepted that some of it is done hormonally. Botanists generally concentrate on ethylene and abscisic acid as culprits in senescence, but neglect gibberellin and brassinosteroid which inhibits root growth if not causing actual root pruning. This is perhaps because roots are below the ground and thus harder to study.

  1. Shoot pruning – it is now known that ethylene induces the shedding of leaves much more than abscisic acid. ABA originally received its name because it was discovered to have a role in leaf abscission. Its role is now seen to be minor and only occurring in special cases.
    • Hormonal shoot pruning theory – a new simple theory says that even though ethylene may be responsible for the final act of leaf shedding, it is ABA and strigolactones that induces senescence in leaves due to a run away positive feedback mechanism. [2] What supposedly happens is that ABA and strigolactones are released by mostly mature leaves under water and or mineral shortages. The ABA and strigolactones act in mature leaf cells however, by pushing out minerals, water, sugar, gases and even the growth hormones auxin and cytokinin (and possibly jasmonic and salicylic acid in addition). This causes even more ABA and strigolactones to be made until the leaf is drained of all nutrients. When conditions get particularly bad in the emptying mature leaf cell, it will experience sugar and oxygen deficiencies and so lead to gibberellin and finally ethylene emanation. When the leaf senses ethylene it knows its time to excise.
  2. Root pruning – the concept that plants prune the roots in the same kind of way as they abscise leaves, is not a well discussed topic among plant scientists, although the phenomena undoubtedly exists. If gibberellin, brassinosteroid and ethylene are known to inhibit root growth it takes just a little imagination to assume they perform the same role as ethylene does in the shoot, that is to prune the roots too.
    • Hormonal root pruning theory – in the new theory just like ethylene, GA, BA and Eth are seen both to be induced by sugar (GA/BA) and oxygen (ETH) shortages (as well as maybe excess levels of carbon dioxide for Eth) in the roots, and to push sugar and oxygen, as well as minerals, water and the growth hormones out of the root cell causing a positive feedback loop resulting in the emptying and death of the root cell. The final death knell for a root might be strigolactone or most probably ABA as these are indicators of substances that should be abundant in the root and if they cannot even support themselves with these nutrients then they should be senesced.
  3. Parallels to cell division – the theory, perhaps even more controversially, asserts that just as both auxin and cytokinin seem to be needed before a plant cell divides, in the same way perhaps ethylene and GA/BA (and ABA and strigolactones) are needed before a cell would senesce.

Seed senescence

Seed germination performance is a major determinant of crop yield. Deterioration of seed quality with age is associated with accumulation of DNA damage. [6] In dry, aging rye seeds, DNA damages occur with loss of viability of embryos. [7] Dry seeds of Vicia faba accumulate DNA damage with time in storage, and undergo DNA repair upon germination. [8] In Arabidopsis , a DNA ligase is employed in repair of DNA single- and double-strand breaks during seed germination and this ligase is an important determinant of seed longevity. [9] In eukaryotes, the cellular repair response to DNA damage is orchestrated, in part, by the DNA damage checkpoint kinase ATM. ATM has a major role in controlling germination of aged seeds by integrating progression through germination with the repair response to DNA damages accumulated during the dry quiescent state. [10]

See also

Related Research Articles

<span class="mw-page-title-main">Germination</span> Process by which an organism grows from a spore or seed

Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ferns, bacteria, and the growth of the pollen tube from the pollen grain of a seed plant.

<span class="mw-page-title-main">Vascular cambium</span> Main growth tissue in the stems, roots of plants

The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xylem inwards, towards the pith, and secondary phloem outwards, towards the bark.

<span class="mw-page-title-main">Plant hormone</span> Chemical compounds that regulate plant growth and development

Plant hormones 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 size, pathogen defense, stress tolerance and reproductive development. Unlike in animals each plant cell is capable of producing hormones. Went and Thimann coined the term "phytohormone" and used it in the title of their 1937 book.

<span class="mw-page-title-main">Auxin</span> Plant hormone

Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essential for plant body development. The Dutch biologist Frits Warmolt Went first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these phytohormones and to determine its chemical structure as indole-3-acetic acid (IAA). Went and Thimann co-authored a book on plant hormones, Phytohormones, in 1937.

<span class="mw-page-title-main">Cytokinin</span> Class of plant hormones promoting cell division

Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and leaf senescence.

Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of plant hormone. It is thought that the selective breeding of crop strains that were deficient in GA synthesis was one of the key drivers of the "green revolution" in the 1960s, a revolution that is credited to have saved over a billion lives worldwide.

<span class="mw-page-title-main">Plant physiology</span> Subdiscipline of botany

Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology, plant ecology, phytochemistry, cell biology, genetics, biophysics and molecular biology.

<span class="mw-page-title-main">Abscisic acid</span> Plant hormone

Abscisic acid 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 stresses, including drought, soil salinity, cold tolerance, freezing tolerance, heat stress and heavy metal ion tolerance.

<span class="mw-page-title-main">Callus (cell biology)</span> Growing mass of unorganized plant parenchyma cells

Plant callus is a growing mass of unorganized plant parenchyma cells. In living plants, callus cells are those cells that cover a plant wound. In biological research and biotechnology callus formation is induced from plant tissue samples (explants) after surface sterilization and plating onto tissue culture medium in vitro. The culture medium is supplemented with plant growth regulators, such as auxin, cytokinin, and gibberellin, to initiate callus formation or somatic embryogenesis. Callus initiation has been described for all major groups of land plants.

<span class="mw-page-title-main">Epidermis (botany)</span> Layer of cells that covers leaves, flowers, roots of plants

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.

Aleurone is a protein found in protein granules of maturing seeds and tubers. The term also describes one of the two major cell types of the endosperm, the aleurone layer. The aleurone layer is the outermost layer of the endosperm, followed by the inner starchy endosperm. This layer of cells is sometimes referred to as the peripheral endosperm. It lies between the pericarp and the hyaline layer of the endosperm. Unlike the cells of the starchy endosperm, aleurone cells remain alive at maturity. The ploidy of the aleurone is (3n) [as a result of double fertilization].

<span class="mw-page-title-main">Gibberellic acid</span> Chemical compound

Gibberellic acid (also called gibberellin A3 or GA3) is a hormone found in plants and fungi. Its chemical formula is C19H22O6. When purified, it is a white to pale-yellow solid.

Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. By contrast, an animal embryo will very early produce all of the body parts that it will ever have in its life. When the animal is born, it has all its body parts and from that point will only grow larger and more mature. However, both plants and animals pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.

<span class="mw-page-title-main">Daphne Osborne</span> British botanist (1930–2006)

Daphne J. Osborne was a British botanist. Her research in the field of plant physiology spanned five decades and resulted in over two hundred papers, twenty of which were published in Nature. Her obituary in The Times described her scientific achievements as "legendary"; that from the Botanical Society of America attributed her success to "her wonderful intellectual style, combined with her proclivity for remarkable and perceptive experimental findings".

<span class="mw-page-title-main">Karrikin</span> A plant growth regulator

Karrikins are a group of plant growth regulators found in the smoke of burning plant material. Karrikins help stimulate seed germination and plant development because they mimic a signaling hormone known as strigolactone. Strigolactones are hormones that help increase growth of symbiotic arbuscular mycorrhizal fungi in the soil, which enhances plant growth and leads to an increase in plant branching.

<span class="mw-page-title-main">Somatic embryogenesis</span> Method to derive a plant or embryo from a single somatic cell

Somatic embryogenesis is an artificial process in which a plant or embryo is derived from a single somatic cell. Somatic embryos are formed from plant cells that are not normally involved in the development of embryos, i.e. ordinary plant tissue. No endosperm or seed coat is formed around a somatic embryo.

Gaseous signaling molecules are gaseous molecules that are either synthesized internally (endogenously) in the organism, tissue or cell or are received by the organism, tissue or cell from outside and that are used to transmit chemical signals which induce certain physiological or biochemical changes in the organism, tissue or cell. The term is applied to, for example, oxygen, carbon dioxide, sulfur dioxide, nitrous oxide, hydrogen cyanide, ammonia, methane, hydrogen, ethylene, etc.

<span class="mw-page-title-main">Strigolactone</span> Group of chemical compounds

Strigolactones are a group of chemical compounds produced by roots of plants. Due to their mechanism of action, these molecules have been classified as plant hormones or phytohormones. So far, strigolactones have been identified to be responsible for three different physiological processes: First, they promote the germination of parasitic organisms that grow in the host plant's roots, such as Strigalutea and other plants of the genus Striga. Second, strigolactones are fundamental for the recognition of the plant by symbiotic fungi, especially arbuscular mycorrhizal fungi, because they establish a mutualistic association with these plants, and provide phosphate and other soil nutrients. Third, strigolactones have been identified as branching inhibition hormones in plants; when present, these compounds prevent excess bud growing in stem terminals, stopping the branching mechanism in plants.

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.

<span class="mw-page-title-main">Ethylene (plant hormone)</span> Alkene gas naturally regulating the plant growth

Ethylene (CH
2
=CH
2
) is an unsaturated hydrocarbon gas (alkene) acting as a naturally occurring plant hormone. It is the simplest alkene gas and is the first gas known to act as hormone. It acts at trace levels throughout the life of the plant by stimulating or regulating the ripening of fruit, the opening of flowers, the abscission (or shedding) of leaves and, in aquatic and semi-aquatic species, promoting the 'escape' from submergence by means of rapid elongation of stems or leaves. This escape response is particularly important in rice farming. Commercial fruit-ripening rooms use "catalytic generators" to make ethylene gas from a liquid supply of ethanol. Typically, a gassing level of 500 to 2,000 ppm is used, for 24 to 48 hours. Care must be taken to control carbon dioxide levels in ripening rooms when gassing, as high temperature ripening (20 °C; 68 °F) has been seen to produce CO2 levels of 10% in 24 hours.

References

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