Plant senescence
Encyclopedia
Plant senescence is the study of aging in plants. It is a heavily studied subject just as it is in the other kingdoms of life. Plants, just like other forms of organisms, seem to have both unintended and programmed aging. Leaf senescence is the cause of autumn leaf color
in deciduous trees.
s. The hormones abscisic acid
and ethylene
are accepted by most scientists as the main causes, but at least one source believes gibberellin
s and brassinosteroids are equally responsible. Cytokinin
s help to maintain the plant cell but when they are withdrawn or if the cell can not receive the cytokinin it may then undergo apoptosis or senescence.
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" 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.
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.
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.
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.
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.
Similarly, auxin and cytokinin may not be enough for plant cell division alone, but a proposed complement of ABA, SA, may be needed in addition.
Autumn leaf color
Autumn leaf color is a phenomenon that affects the normally green leaves of many deciduous trees and shrubs by which they take on, during a few weeks in the autumn season, one or many colors that range from red to yellow...
in deciduous trees.
Programmed senescence
Programmed senscence seems to be heavily influenced by plant hormonePlant hormone
Plant hormones are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and, when moved...
s. The hormones 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:...
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...
are accepted by most scientists as the main causes, but at least one source believes gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....
s and brassinosteroids are equally responsible. Cytokinin
Cytokinin
Cytokinins are a class of plant growth substances 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...
s help to maintain the plant cell but when they are withdrawn or if the cell can not receive the cytokinin it may then undergo apoptosis or senescence.
Annual versus perennial benefits - theory
Some plants have evolved into annualsAnnual plant
An annual plant is a plant that usually germinates, flowers, and dies in a year or season. True annuals will only live longer than a year if they are prevented from setting seed...
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
Perennial plant
A perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...
. 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
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...
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
Head start (positioning)
In positioning, a head start is a start in advance of the starting position of others in competition, or simply toward the finish line or desired outcome...
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.
Plant self pruning - theory
There is a speculative hypothesisHypothesis
A hypothesis is a proposed explanation for a phenomenon. The term derives from the Greek, ὑποτιθέναι – hypotithenai meaning "to put under" or "to suppose". For a hypothesis to be put forward as a scientific hypothesis, the scientific method requires that one can test it...
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 meristemMeristemA meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
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.
Hormonal induction of senescence - theory
There is not a lot of 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 ethyleneEthylene
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...
and 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:...
as culprits in senescence, but neglect gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....
and brassinosteroid
Brassinosteroid
Brassinosteroids are a class of polyhydroxysteroids that have been recognized as a sixth class ofplant hormones. These were first explored nearly forty years ago when Mitchell et al. reported promotion in stem elongation and cell division by the treatment of organic extracts of rapeseed pollen...
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 ethylene induces senescence in leaves due to a run away positive feedback mechanism. What supposedly happens is that ethylene is released by mostly mature leaves under water and or mineral shortages. The ethylene acts in mature leaf cells however, by pushing out minerals, water, sugar, gases and even the growth hormones auxinAuxinAuxins 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...
and cytokininCytokininCytokinins are a class of plant growth substances 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...
(and possibly salicylic acidSalicylic acidSalicylic acid is a monohydroxybenzoic acid, a type of phenolic acid and a beta hydroxy acid. This colorless crystalline organic acid is widely used in organic synthesis and functions as a plant hormone. It is derived from the metabolism of salicin...
in addition). This causes even more ethylene to be made until the leaf is drained of all nutrients.
- Hormonal shoot pruning theory - a new simple theory says that ethylene induces senescence in leaves due to a run away positive feedback mechanism. What supposedly happens is that ethylene is released by mostly mature leaves under water and or mineral shortages. The ethylene acts in mature leaf cells however, by pushing out minerals, water, sugar, gases and even the growth hormones auxin
- 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 and brassinosteroid 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 are seen both to be induced by sugar and gas shortages in the roots, and to push sugar and gases, as well as minerals, water and the growth hormones out of the root cell causing a positive feedback loop resulting the emptying and death of the root cell.
- 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 are needed before a cell would senesce.
- Discussion of the complete mechanism - what really may occur is that because ethylene pushes out all nutrients out of the shoot cell including sugar and gases, eventually this causes a shortage of these nutrients in the organ (the shoot) which is supposed to procure them. This shortage leads to GA/BA synthesis in the rapidly declining shoot cell, and this simply adds fuel to the fire.
- A role for abscisic acid - finally a question may be to ask, what is the role of abscisic acid, the hormone which was first thought to be the primary mover in this department? According to author of this theory, ABA is induced when plant cells are encountering stress other than that of a nutrient shortage kind. In this case a senescing cell experiencing a drain of nutrients may experience a strain which causes it to produce ABA. Indeed it may not be that ethylene and GA/BA alone are needed for programmed cell senescence, but that all three are needed.
Similarly, auxin and cytokinin may not be enough for plant cell division alone, but a proposed complement of ABA, SA, may be needed in addition.