Plant senescence is the process of aging in plants. Plants have both stress-induced and age-related developmental aging. Chlorophyll degradation during leaf senescence reveals the

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

s, such as anthocyanin and xanthophylls, which are the cause of

autumn leaf color Autumn leaf color is a phenomenon that affects the normal green leaves of many deciduous trees and shrubs by which they take on, during a few weeks in the autumn season, various shades of yellow, orange, red, purple, and brown. The phenomenon i ...

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.

Hormonal regulation of senescence

Programmed senescence seems to be heavily influenced by

plant hormone 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 ...

s. The hormones

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 ...

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 ...

jasmonic acid Jasmonic acid (JA) is an organic compound found in several plants including jasmine. The molecule is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway. It was first isolate ...

and

salicylic acid Salicylic acid is an organic compound with the formula HOC6H4CO2H. A colorless, bitter-tasting solid, it is a precursor to and a metabolite of aspirin (acetylsalicylic acid). It is a plant hormone, and has been listed by the EPA Toxic Substa ...

are accepted by most scientists as promoters of senescence, but at least one source lists

gibberellin 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 plan ...

s, brassinosteroids and strigolactone as also being involved.

Cytokinin 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 lea ...

s help to maintain the plant cell and expression of cytokinin biosynthesis genes late in development prevents leaf senescence. A withdrawal of or inability of the cell to perceive cytokinin may cause it to undergo

apoptosis Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes ( morphology) and death. These changes in ...

or senescence. In addition, mutants that cannot perceive

show delayed senescence. Genome-wide comparison of

mRNAs In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of synthesizing a protein. mRNA is created during the p ...

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.

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 A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...

. This may be a programmed "strategy" 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 A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...

production rather than saving some for the plant to overwinter, which would limit seed production. 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

, then there will be a far lesser chance that a random mutation will slow the bug compared to more diverse annuals.

Plant self-pruning

There is a speculative

hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...

on how and why a plant induces part of itself to die off. 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. * Poor productivity reasons for plant self pruning – the plant rarely prunes young dividing

meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...

atic cells, but if a fully grown mature cell is no longer acquiring nutrients that it should acquire, then it is pruned. ** Shoot efficiency self pruning reasons – for instance, presumably a mature shoot cell must on average produce enough sugar, and acquire enough oxygen and carbon dioxide to support both it and a similar sized root cell. Actually, since plants are obviously interested in growing it is arguable, that the "directive" of the average shoot cell, is to "show a profit" and produce or acquire more than enough sugar and gases than is necessary to support both it and a similar sized root cell. If this "profit" isn't shown, the shoot cell is killed off and resources are redistributed to "promising" other young shoots or leaves in the hope that they will be more productive. ** Root efficiency self pruning reasons – similarly a mature root cell must acquire on average, more than enough minerals and water needed to support both it and a similar sized shoot cell that does not acquire water and minerals. If this does not happen, the root is killed off and resources sent to new young root candidates. * Shortage/need-based reason for plant self pruning – this is the other side of efficiency problems. **Shoot shortages – if a shoot is not getting enough root derived minerals and water, the idea is that it will kill part of itself off, and send the resources to the root to make more roots. **Root shortages – the idea here is that if the root is not getting enough shoot derived sugar and gases it will kill part of itself off and send resources to the shoot, to allow more shoot growth. 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. Plant scientists generally concentrate on

and

as culprits in senescence, but neglect

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. #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 Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop which exacerbates the effects of a small disturbance. That is, the effects of a perturbation on a system include an increase in th ...

mechanism. 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 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 ...

and

cytokinin 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 lea ...

(and possibly jasmonic and

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 Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or do ...

and

oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...

deficiencies and so lead to

and finally ethylene emanation. When the leaf senses ethylene it knows its time to excise. #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

, brassinosteroid and

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 Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...

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. #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 A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...

germination 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, fe ...

performance is a major determinant of

crop yield In agriculture, the yield is a measurement of the amount of a crop grown, or product such as wool, meat or milk produced, per unit area of land. The seed ratio is another way of calculating yields. Innovations, such as the use of fertilizer, the c ...

. Deterioration of seed quality with age is associated with accumulation of

DNA damage DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA d ...

. In dry, aging rye seeds, DNA damages occur with loss of viability of

embryos An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

. Dry seeds of ''

Vicia faba ''Vicia faba'', commonly known as the broad bean, fava bean, or faba bean, is a species of vetch, a flowering plant in the pea and bean family Fabaceae. It is widely cultivated as a crop for human consumption, and also as a cover crop. Varieti ...

'' accumulate DNA damage with time in storage, and undergo

DNA repair DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA d ...

upon

. In ''

Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organ ...

'', a

DNA ligase DNA ligase is a specific type of enzyme, a ligase, () that facilitates the joining of DNA strands together by catalyzing the formation of a phosphodiester bond. It plays a role in repairing single-strand breaks in duplex DNA in living orga ...

is employed in repair of DNA single- and double-strand breaks during seed germination and this ligase is an important determinant of seed longevity. In

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...

s, 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.

References

*Special issue about plant senescence i
Plant Biology volume 10 issue s1

External links

* ttp://www.planthormones.info/ The Start at a General Theory of Plant Senescence {{senescence Plant physiology Senescence in non-human organisms