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Auxins are a class of
plant growth substance (often called
phytohormones or plant hormones). Auxins
play an essential role
in coordination of many growth and
behavioral processes in the plant life
cycle.
Auxins, the first
hormones to be discovered, have been
demonstrated to be a basic coordinative
signal of plant development. Their
pattern of active transport through the
plant is complex, and auxins typically
act in concert with (or opposition to)
other plant hormones. For example, the
ratio of auxin to cytokinin in certain
plant tissues determines initiation of
root versus shoot buds. As a result, a
plant can (as a whole) react on external
conditions and adjust to them, without
requiring a nervous system.
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Auxins directly
stimulate or inhibit the expression of
specific genes. Auxin induces
transcription by targeting for
degradation members of the Aux/IAA
family of transcriptional repressor
proteins, The degradation of the
Aux/IAAs leads to the derepression of
ARF-mediated transcription. Aux/IAAs are
targeted for degradation by
ubiquitination, catalysed by an SCF-type
ubiquitin-protein ligase.
In 2005, it was
demonstrated that the F-box protein
TIR1, which is part of the ubiquitin
ligase complex SCFTIR1, is an auxin
receptor. Upon auxin binding TIR1
recruits specific transcriptional
repressors (the Aux/IAA repressors) for
ubiquitination by the SCF complex. This
marking process leads to the degradation
of the repressors by the proteasome,
alleviating repression and leading to
specific gene expression in response to
auxins.
Another protein
called ABP1 (Auxin Binding Protein 1) is
a putative receptor, but its role is
unclear. Electrophysiological
experiments with protoplasts and
anti-ABP1 antibodies suggest that ABP1
may have a function at the plasma
membrane.
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On the cellular
level, auxin is essential for cell
growth, affecting both cell division and
cellular expansion. Depending on the
specific tissue, auxin may promote axial
elongation (as in shoots), lateral
expansion (as in root swelling), or
isodiametric expansion (as in fruit
growth). In some cases (coleoptile
growth) auxin-promoted cellular
expansion occurs in the absence of cell
division. In other cases, auxin-promoted
cell division and cell expansion may be
closely sequenced within the same tissue
(root initiation, fruit growth). In a
living plant it appears that that auxins
and other plant hormones nearly always
interact to determine patterns of plant
development.
According to the
"acid growth theory," auxins may
directly stimulate the early phases of
cell elongation by causing responsive
cells to actively transport hydrogen
ions out of the cell, thus lowering the
pH around cells. This acidification of
the cell wall region activates enzymes
known as expansins, which break bonds in
the cell wall structure, making the cell
wall less rigid. When the cell wall is
partially degraded by the action of
auxins, this now-less-rigid wall is
expanded by the pressure coming from
within the cell, especially by growing
vacuoles.
However, the acid
growth theory does not by itself account
for the increased synthesis and
transport of cell wall precursors and
secretory activity in the Golgi system
that accompany and sustain
auxin-promoted cell expansion.
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Growth and division
of plant cells together result in growth
of tissue, and specific tissue growth
contributes to the development of plant
organs. Growth of cells contributes to
the plant's size, but uneven localized
growth produces bending, turning and
directionalization of organs, for
example, stems turning toward light
sources (Phototropism), growth of roots
in response to gravity (gravitropism),
and other tropisms.
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