Plant Physiology 2008

"How plants grow": Lecture on auxin, the primary growth hormone of plants.

I. What is growth?

for both animal and plants:

growth = irreversible increase in volume or size (of an organism)

Plants are radically different then animals when it comes to growth and development

animals: determinate growth - animals have a pre-set size and shape. animals grow by adding more
cells to an already established part of the organism (we grow in size because more cells of all kinds
are added in proportion to say a leg or arm).

plants: indeterminate growth - plants do not have a pre-set size and final shape.......there is no
defined “end” volume or size. Plants grow by increasing the size of their cells once a cell is formed in
a tissue;

growth can occur in plants without cell division!

II. Plants induce their cells to grow by secreting auxin from the apical meristems and
secondary meristems

Auxin, (or IAA), is one of 5 major plant hormones. When cells are exposed to auxin in within the plant,
the increase in volume by as much as 20 times their original size.

What is a hormone?

The classic concept of what a hormone is was established in animal systems:

-a molecule that is produced in one part of body

-transported to another part of the body
-elicits a specific response
-effective at minute amounts

..........example: insulin.........produced in pancreas, secreted into blood, travels to distant tissues to

stimulate glucose uptake
Auxin fits this definition:

-produced in apical meristem cells

-transported to cells distant from meristem
-elicits a growth response
-effective in minute amounts

A. Auxin

1. site of auxin synthesis- apical meristems (root and shoot) and secondary meristems (e.g., leaf
meristem)........cause of phenomenon of apical dominance

a. What is an apical meristem?

A compact lens shaped cluster of undifferentiated mother cells located at the very tips of the shoots
and roots. Also in vegetative "buds" in growing shoots and axillary branches. When these "stem"
cells divide, the daughter cells undergo differentiation and maturation to form the plant body.

Auxin is secreted by the apical meristems and flows basipetally via the apoplast to the
cells beneath

Auxin Auxin

Auxin Auxin

2. site of auxin action - auxin elicits a growth response in cells and tissues located basipetally
from the apical meristems (the basipetal means the cells and tissue beneath the apical meristems)
3. mechanism of auxin action: “acid induced growth model”

Auxi Auxin
n Cell in growing zone

The overall mechanism of how auxin induces growth of cells via increasing the size of the cell is mediated by
(i) auxins immediate activation/stimulation of H+/ATPases (proton pumps) on the plasma membranes and (ii)
auxin activation of growth genes in the nucleus.

The step-by-step mechanism of how this occurs is as follows:

1. As auxin the seeps into the cell wall, it contacts and binds a special auxin cell membrane protein receptor
that transports the hormone into the cell.

2. Auxin immediately activates the H+/ATPases (proton pumps) on the plasma membranes.......
this causes the cell wall pH to decrease from a neutral pH 7 to and acid pH 5

3. Enzymes called cellulose hydrolases in the cell wall are activated by the acidic pH ..........hydrolase enzymes
break the weak hydrogen bonds that cross-link linear cellulose molecules together in a cellulose microfibril
4. When these H-bonds are severed, the cellulose strands in the cell wall become “loose” or “slippery” and
yield to the cells turgor pressure

5. the H+/pumps are also electrogenic........that is to say they hyperpolarize the membrane so that it is even
more (-) inside the cell and than outside the cell in the cell wall

6. K+ ions that are normally held in the cell wall flow into the cell via K+ channels in response to the negative
charge gradient

7. Because K+ is an osmotic particle (and H+ ions are not), increased amounts of K+ in the cell cause a more
negative cell Ψw, so H2O "gushes" into the cell further increasing the turgor pressure

8. the loosened cell wall responds by slipping........the cell wall expands or "stretches" when this happens,
although it makes it thinner

9. the cell then expands..........it elongates......it increases in volume......this is growth

10. collectively, all the cells in this tissue are simultaneously expanding thus making that part of the plant
increase in size or length.

But to sustain the growth, the cell now has to make more cell wall and more cell membrane:

-the cell wall cannot be allowed to get too thin

-and the plasma membrane cannot not stretch or slip like the cell wall

Auxin also acts to sustain the acid induced growth by turning on the genes required for synthesis:

of new cell wall materials : pectins, cellulose, lignin’s

of new cell membrane materials: phospholipids, membrane proteins

To do this, that is to turn on genes, it takes at least one to several hours

-once the genes are turned on, growth can be sustained for days until the meristem is separated from the
growing section of cells
Thus, the mode of auxin action is often considered a 2 phase response:

1. fast mode (minutes) : turn on plasma membrane H+ pumps.......growth is initiated

2. slow mode (hours) : turn on biosynthetic genes...........................growth is sustained

a. Weed-be-gone herbicide (a.k.a. 2,4-D) - kills weeds by causing them to grow-to-death!

2,4-D is a synthetic auxin.......resembles IAA, the naturally occurring plant hormone

Mechanism of action of 2,4-D herbicide: 2,4-D triggers the fast-mode auxin response but it cannot also
trigger the slow-mode auxin response (biosynthesis of new cell membrane/cell wall).

The herbicide kills weeds by causing:

rapid elongation of cells

cell walls stretch

becoming too thin

no new cell wall or

membrane is made
so the cell wall
becomes dangerously
thin as the cell
elongates

the cell finally bursts and

dies due to the inability of
the thin cell wall to contain
the enormous internal cell
water pressure