LECTURE 17 - Root Shoot Balance, Apical dominance

Apical Dominance
Typically a shoot has leaves attached to a stem. The apex of the stem has an
actively growing apical bud. It produces additional nodes and internodes to add to
the length of the shoot. Additional leaves appear at the nodes as they are produced.
In some plants, the lateral bud located in the axil of each leaf does not grow to
form branches, especially at first. This condition is known as strong apical
dominance.

Apical dominance is thought to be caused by the apical bud producing IAA (auxin)
in abundance. This auxin is transported basipetally (of or relating to the
development or maturation of tissues or organs or the movement of substances)
from the apical bud. The auxin causes the lateral buds to remain dormant.


Apical dominance occurs when the shoot apex prevents the growth of lateral buds
so that the plant may grow vertically. It is important for the plant to devote energy
to growing upward so that it can get more light to undergo photosynthesis. If the
plant utilizes available energy for growing upward, it may be able to outcompete
other individuals in the near vicinity. Plants that were capable of outcompeting
neighboring plants likely had higher fitness. Apical dominance is therefore most
likely adaptive.

Typically, the end of a shoot contains an apical bud, which is the location where
shoot growth occurs. The apical bud produces an auxin (IAA) that inhibits growth
of the lateral buds further down on the stem. It was first discovered that the plant
hormone auxin likely regulates apical dominance in 1934.

Auxin is predominantly
produced in the growing shoot apex and is transported throughout the plant via the
xylem and diffuses into lateral buds which prevent elongation.

When the apical bud is removed, the lowered IAA concentration allows the lateral
buds to grow and produce new shoots, which compete to become the lead growth.
Root: Shoot Balance

Nitrogen up-take and use in a tree have been examined using shoot/root models.
The most tested and effective is called the Thornley model. To recognize resource
allocation patterns in trees between shoot and root (approaching a functional
balance), only four components are required: sapwood shoot mass, sapwood root
mass, photosynthesis rate, and nitrogen uptake rate. The model equation is shown
in Figure 1.

Figure 1:

Trees attempt to balance shoot mass and photosynthesis rates against root mass and
nitrogen uptake. A tree will adjust the living mass of roots or shoots to correct any
deficiency in photosynthesis rates or nitrogen uptake. Carbohydrate shortages /
nitrogen increases will initiate more shoots--nitrogen shortages / carbohydrate
increases will initiate more roots.

Both of the benchmark processes (and associated tissue mass) listed above, and as
shown in Figure 1, must always be functionally balanced across the tree. For
example, as nitrogen absorption declines, what nitrogen remains is concentrated
more in the roots and used preferentially. This leads to less shoot growth and more
root growth. Even before growth is noticeably reduced, the tree is reallocating
nitrogen to vital processes. One vital need is in the absorbing roots where more
rapid turn-over is occurring as nitrogen concentrations fall.

With supplemental nitrogen fertilizers, root growth declines and shoot growth
increases. In addition, the added nitrogen causes a decline in starch and an increase
in sugars. Increased sugar contents and additional nitrogen availability generate
improved access and attack conditions for a number of pests.

Q What factors affect root-shoot ratios?

The ratio of shoot to root growth varies widely between species, changes between
phases in plant development, and can be modified by external conditions.

At a gross level the ratio between shoot and root weight is a generally
characteristic of the family or group.

If either roots or shoots are pruned, the plant will tend to recover to the
characteristic ratio.

If external conditions change, shoot-root ratio will be affected.

The root-shoot ratio increases if water is withheld from the rooting medium (Sharp
and Davies, 1979)