Compare leaf senescence, fruit ripening and abscission with each other.

Leaf senescence is a process in deciduous trees that allows reabsorption of nutrients.

Senescence is followed by abscission, where dead and obsolete leaves are dropped. Fruit
ripening increases the palatability of the fruit, increasing its attractiveness to animal seed
disseminators. The functions of these three processes underlie their greatest differences: the
purpose of the colour changes in senescence and ripening, and the presence of the
climacteric in fruit ripening only. However, the three processes share several underlying
mechanisms, including the role of ethylene as a positive feedback regulator, auin as an
inhibitor of onset, and changing sensitivity to ethylene as the process accelerates.
Leaf senescence, fruit ripening and abscission all serve a different purpose to the plant.
!bout "#$ of the % and & from senescing leaves is reabsorbed, which is a ma'or source of
nutrients for growth in the net growing season. (n one eperiment, shrubby oaks were
defoliated late in the season, preventing retranslocation of nutrients. (n the subse)uent year
they showed a large decrease in acorn production, compared to controls and non*defoliated
shaded plants. !bscission can involve drop of senesced leaves to conserve water over
winter, or dropping unnecessary tissues such as flower petals post*fertilisation. +ropping a
proportion of fruits before they ripen can ensure that the plant has enough resources to bring
the remaining fruits to maturity. The purpose of fruit ripening is to attract animals to
disseminate the seeds of the plant , plants therefore make their fruits appealing by
increasing the concentrations of soluble sugars and volatilising secondary compounds. The
secondary metabolite capsaicin, found in chillies, has a highly aversive taste to mammals
but is tolerated by birds. This may be a strategy to ensure wide*range dispersal of the chilli
seeds, deterring mammalian dispersers who may not travel as far as birds.
Leaf senescence and fruit ripening both involve colour changes. (n leaf senescence,
photosystem (( breaks down. -hlorophyll b reductase reduces chlorophyll b to chlorophyll a,
which destabilises &S((. -hlorophylls subse)uently degrade into colourless tetrapyrroles, so
that other pigments, like anthophylls and carotenes, are revealed. !nthocyanins are
synthesised de novo once around half of the chlorophyll has been degraded. This is thought
to be a means of overcoming photoinhibition. &hotoinhibition can result in reduced
reabsorption of foliar nutrients, which can decrease tree fitness, as seen in the above
eample with shrubby oaks. (n fruit ripening, chlorophyllases are induced to destroy
chlorophylls. -hlorophyllases are also involved in de*greening in leaf senescence. Like
anthocyanin in leaf senescence, the pigment lycopene is synthesised de novo in ripening
tomato fruits. However, fruit ripening is also associated with a conversion in plastid type:
chloroplasts are converted into chromoplasts, which contain pigments.
There is a rise in -./ production, uni)ue to fruit ripening, known as the climacteric. 0hile
1%! and protein concentration in fruits remains approimately the same throughout
ripening, the synthetic capacity for both 2measured by ability of plants to incorporate labelled
precursors3, is enhanced during the early part of the climacteric rise, and then declines.
-ycloheimide, an inhibitor of eukaryotic protein synthesis, only inhibits ripening during the
early part of the climacteric, after which all essential en4ymes have been synthesised. The
climacteric is not due to protein synthesis: cycloheimide inhibits ripening without affecting
the climacteric. This follows from the fact that the abscission 4one doesn5t show a climacteric
during abscission, despite an increase in protein synthesis 2of cell wall degrading en4ymes ,
cellulases, pectinases and peroidases3. The climacteric is more likely associated with
thermogenesis, an increase in heat in ripening fruits, the purpose of which is to volatilise
secondary compounds to attract animal seed disseminators. !lternative oidase redirects
electrons from ubi)uinone in the respiratory chain into a non*phosphorylating pathway,
dissipating the energy as heat. (n mangoes, the alternative oidase is undetectable in unripe
fruits, and increases during ripening.
6thylene positive feedback loops are involved in fruit ripening, leaf senescence, and
abscission. &ositive feedback loops are ubi)uitous in biology, and in physiology are usually
associated with a climactic event. They ensure that this event occurs, as the path to it is self*
reinforcing. (n climacteric fruits, ripening upregulates ethylene production, and ethylene
upregulates ripening. 6thylene is produced by senescing leaves and increases the
progression of senescence. The abscission 4one also produces ethylene, which regulates
abscission.
%egative feedback ensures that senescence, ripening or abscission are not prematurely
triggered. (n the early stages of fruit ripening, tissues show autoinhibition in response to
ethylene. This protects the fruit from premature ripening in response to the ethylene bursts
accompanying anthesis and pollination. (ncreased sensitivity to ethylene can induce
autocatalysis. Sensitivity is induced by low auin levels , auin inhibits onset of fruit ripening,
and applying &-(7!, an auin analogue which triggers destruction of auin, brings forward
the onset of ripening. !uin also delays the onset of senescence in leaves. .ther hormones
can also delay senescence, and the choice of hormone is adaptive , for eample,
dandelions use cytokinin in the summer to delay leaf senescence, as it is beneficial to
control leaf life as a function of soil fertility, etending the leaf5s use to the plant for as long as
possible. !uin inhibits early leaf abscission: applying auin to the cut ends of eplants
delays abscission in fresh eplants, but accelerates abscission in older eplants. 6thylene
has no effect on fresh eplants and accelerates abscission in older material. From these
results it was concluded that there are two stages of leaf abscission: stage (, before the
tissue is ethylene responsive, and stage ((, where ethylene triggers abscission. Sensitivity to
ethylene may play a role in abscission as it does in the ripening of climacteric fruits: the
tomato homologue of the ethylene receptor that mediates seedling responses in Arabidopsis
is only epressed at detectable levels in abscission 4ones that are autocatalytically
synthesising ethylene.
(n conclusion, several of the differences between leaf senescence, abscission and fruit
ripening are reflected by the differing functions of the processes. The colour change in
ripening fruits is associated with increasing their attraction to animal seed disseminators,
whereas colour changes in senescing leaves are due to reabsorption of % and & from
pigments in the leaf, and protection of the leaf from photoinhibition. The presence of the
climacteric in ripening fruits but not in abscising leaves is similarly an adaptation to increase
attractiveness to animal seed disseminators via volatilisation of secondary compounds.
However, the underlying mechanisms of all three processes have overarching similarities.
6thylene is a positive regulator of senescence, abscission and ripening, and ethylene is
produced in senescing and ripening tissues, and in the abscission 4one. This results in a
positive feedback loop. &ositive feedback in abscission and 2in some climacteric fruits3 fruit
ripening is only allowed once tissues have increased sensitivity to ethylene, which in tomato
at least is due to increased epression of an ethylene receptor. %egative feedback from
other hormones , auin is a negative regulator in all three processes , can delay onset, and
the choice of hormone controlling senescence in particular is adaptive, depending on the
growth strategy of the plant.