Auxin Functions

One of the main functions of auxins in a plant is cell elongation. In the stems, a steady supply of
auxin is needed for the elongation in the subapical regions of the plants. The optimal concentration for
elongation in the stem is 10-6 to 10-5 M; some plants, like Arabidopsis, may have a lower optimal auxin
concentration. In figure x, it can be seen that in concentrations higher than the optimal, auxin becomes
inhibitory as it induces ethylene biosynthesis, which inhibits growth. In the root, only relatively lower
concentrations of auxin, around 10 -10 to 10-9 M, promotes the growth of intact roots. Higher
concentrations, around the concentrations that induces growth in the stem, would inhibit root elongation
(Taiz & Zeiger, 2010).

Figure x. This is a typical dose-response curve

for auxin-induced growth in pea stems or oat
coleoptiles (Taiz & Zaiger, 2010).
Cell elongation and expansion in plants take place in three steps: osmotic uptake of water driven
by a water potential, turgor pressure buildup due to rigidity of the cell wall, and biochemical cell wall
loosening for expansion. Auxin is said to induce the loosening of the plant cell wall in a mechanism
described as the acid growth hypothesis, in which hydrogen ions acts as the intermediate. The activity of
the H+-ATPase in the plasma membrane is increased by the presence of auxin. The hypothesis predicts
five main things: only acid buffers can produce the desired response, auxin should increase the rate of
wall acidification and the kinetics of wall acidification should match the auxin-induced growth, neutral
buffers should stop growth caused by auxins, other substances that promote wall acidification should
induce growth, and the cell wall loosening factor should have optimal activity at an acidic pH. These
predictions have been proven through various experiments. As seen in figure xx, auxin-stimulated wall
acidification is consistent with the growth kinetics of the plant. Expansins, which loosen the cell walls of

plant cells, only break the hydrogen bonds of the polysaccharides in the cell well when the pH is acidic
(Taiz & Zeiger, 2010).

Figure xx. The kinetics of the cell elongation

matches the lowering of the pH caused by the
presence of auxin (Jacobs and Ray, 1976).
Auxin is also observed to be mostly affecting the outer tissues of a plant stem. When a stem is cut
lengthwise and grown in a solution without auxin, the inner vascular tissues grow faster, resulting in
stems bending outward. However, when it is grown in an auxin solution, the outer tissues, like the
epidermis and the cortex, now grow faster than the inner tissues, resulting in the stems growing inward.
This proves that auxin primarily targets the outer tissues in its action in the plant. This is especially
evident in the auxin-mediated phototropic response of a plant. When a plant is given a directional light
stimulus, the auxin produced at the apex is transported laterally towards the shaded side before going
down the stem. Two flavoproteins, phototropins 1 and 2, are said to be the photoreceptors that induces the
phototropic response. When stimulated by blue light, the phototropins autophosphorylate and display a
lateral gradient in the stem. This results in the dissociation of the phototropins from the plasma
membrane, making it free for interactions with auxin transporters and its regulators. The lateral gradient
of phototropins seem to induce the movement of the auxin towards the shaded side of the stem. When it
reaches its target area, auxin only promotes the growth of one side of the stem, producing a differential
growth (Figure xx) towards the source of light (Taiz & Zaiger, 2010).

Figure xxx. The growth response of the shaded and

irradiated sides of the plant shows differential growth
(Iino & Briggs, 1984).
Auxin also has roles in the gravitropic response of plants. Early experiments have shown that a
horizontally oriented coleoptile will redistribute its auxin to the lower side, increasing that sides growth,
and curving upwards. It is hypothesized that the plants detect gravity via large and dense amyloplasts,
which can sediment at the bottom of the cell. Such amlyoplasts are called statoliths, with the cells
containing them called statocytes. In the shoots and coleoptiles, a starch sheath surrounding the vascular
tissues perceives gravity. On the other hand, gravity is detected by the roots in the columella, or the root
cap. The current hypothesis on how statocytes sense the falling statoliths is called the starch-statolith
hypothesis. Presumably, the pressure resulting from the statolith on the endoplasmic reticulum triggers the
response (Taiz & Zeiger, 2010).
Secondary messengers are thought to conduct the information to various parts of the plant for
corrective differential growth due to a misalignment with the gravity vector. Ca 2+ and pH gradients are the
most likely candidates as secondary messengers for gravistimulation. In the columella, the pH changes
can be observed in the cells responding to gravity. It is hypothesized that the activation of H +-ATPase is
one of the initial steps in gravity perception, resulting in the directional uptake and efflux of auxin in the

affected cells. On the other hand, when Ca 2+ is removed from the roots through chelators inhibits the
gravitropic response of the roots (Taiz & Zaiger, 2010).
Like in apex, auxin can also be distributed laterally in the root cap due to changes in the direction
of gravity. When root is oriented vertically, auxin transport is equal on all sides. However, when the root
is positioned horizontally, the root cap directs most of the auxin to the lower side. As mentioned above,
high concentrations of auxin the root causes inhibition of growth, thus the lower sides elongation would
be slowed (Taiz & Zeiger, 2010).
Auxin also influences the development of the plant. Indeed, the morphology of the plant depends
on movement of the auxin using the polar transport system, maintaining the basic shoot-root polarity and
the polar outgrowths. One of its developmental effects includes the regulation of apical dominance. When
the apex is removed, lateral buds are usually seen growing. However, when auxin is applied at the
decapitated apex, this is prevented. The direct inhibition model suggests that the optimal auxin
concentration for bud growth is much lower than those found the stem; as such, normal auxin
concentrations tend to inhibit the growth of the bud. If this is the case, auxin concentration should be
lower in the axillary buds when the apex is removed (Taiz & Zeiger, 2010). However, a study by
Langridge and colleagues (1989) showed that the reverse is true. The current hypothesis is that other
hormones, like cytokinins and abscisic acid, may work together with auxin to bring about apical
dominance in plants.
The appearance of lateral and adventitious roots in plants seems to be also regulated by auxin.
High auxin concentrations in the root promote the division of pericycle cells in the areas above the root
elongation zones, producing lateral roots. On the other hand, the formation of adventitious roots is
somewhat parallel to the formation of lateral roots. Clusters of mature cells in non-root tissue locations
that renew their cell division activity become root apical meristems and form these adventitious roots
(Taiz & Zeiger, 2010).
The presence of auxin can also delay the proliferation of abscission layers in leaves preceeding
senescence. When leaves, flowers or fruits are set to be shed, an abscission zone, containing differentiated
layer of cells called the abscission layer, forms. This can only be seen in older leaves, in which auxin
levels are definitely lower than when they were younger. When leaves are excised from a plant, an
abscission layer quickly forms in the petiole. However, when auxin is applied to the petiole, the formation
of the abscission layer is inhibited (Taiz & Zeiger, 2010).
A study by Kuhlemeier and Reinhardt (2001) treated Arabidopsis plants with an auxin transport
inhibitor. The inhibition of auxin caused abnormal floral development. This implies that the polar
transport of auxin in the inflorescence meristem is vital in the development of flowers. This also holds
true in the formation of normal phyllotaxy in the leaves of the plant.

Auxin is also present in the pollen, in the endosperm, and in the embryo of developing seeds,
suggesting that it has a role in the development of the fruit. It is suggested that after fertilization, the
growth of the fruit may depend on the auxin produced by the seeds. At initial stages of fruit growth, the
endosperm may be the primary source of auxin in the fruit. At later stages, the growing embryo may take
over as the source of auxin for fruit development (Taiz & Zeiger, 2010).
A recent study discovered that the role of auxin goes even further back from fruit development.
Auxin is also responsible in the regulation of the development of the female gametophyte in Arabidopsis.
The seven-cell structure, containing an egg cell, a central cell, two synergid cells and three antipodal cells,
is influenced by auxin gradients within the embryo sac. The proper formation of the specific pattern of
cells within the embryo sac requires auxin distribution and signaling (Pagnussat et al., 2009).
REFERENCES
Iino, M., & Briggs, W. R. (1984) Growth distribution during first positive phototropic curvature of maize
coleoptiles. Plant Cell Environment, 7: 97104.
Jacobs, M, & Ray, P. M. (1976) Rapid auxin-induced decrease in the free space pH and its relationship to
auxin-induced growth in maize and pea. Plant Physiology, 58: 203209.
Kuhlemeier, C. & Reinhardt, D. (2001) Auxin and Phyllotaxis. Trends in Plant Science. 6: 187189.
Langridge, W. H. R., Fitzgerald, K. J., Koncz, C., Schell, J., & Szalay, A. A. (1989) Dual promoter of
Agrobacterium tumefaciens mannopine synthase genes is regulated by plant growth hormones.
Proceedings of the National Academy of Sciences USA, 86: 32193223.
Pagnussat GC, Alandete-Saez M, Bowman JL, & Sundaresan V. (2009). Auxin-dependent patterning and
gamete specification in the Arabidopsis female gametophyte. Science, 324:16849.
Taiz, L. & Zeiger, E. (2010). Plant Physiology, Fifth Edition. Sunderland, Massachussets, USA: Sinauer
Associates, Inc. Publishers.