1.

10AQUAPORINS FACILITATE

THE CELLULAR MOVEMENT

OF WATER

Porinsare a class of membrane proteins that belong

to a large family of proteins called major intrinsic

proteins (MIPs)that are found in the cell membranes

of all living organisms including plants, microorganisms, and animals. Porin-type channels are
nonselective

cation channels which are characterized by aβ-pleated

sheet protein structure. In plants, porins are generally restricted to the outer membranes of
mitochondria

and chloroplasts and account for their highly permeable properties. In contrast,aquaporinsare
membrane

protein channels or pores controlling the selective movement of water primarily. This is indicated by
the fact

that the presence of aquaporins does not affect the

electrical conductance of a membrane which indicates

that small ions such as H

are not conducted by these

membrane channels. The results of genome sequencing

indicate that Arabidopsis thalianaexhibits 35 aquaporin

gene homologs and 33 homologues have been detected

in the rice genome. Thus, plants exhibit a multiplicity of

aquaporin isoforms which, in part, reflects the multiplicity of internal membrane types in which
these channels

are localized within a plant cell (Figure 1.10A). For

example, 13 homologues have been detected in plant

plasma membranes of Arabidopsis and are designated

PIPs forplasma membrane intrinsicproteins whereas

the tonoplast membrane, which surrounds the inner

vacuole of plant cells, exhibits 10 homologues and are

designated TIPs.
The structure of aquaporins is highly conserved

between plants, microbes, and animals. Four subunits

(tetramers) associate to form a single plant aquaporin.

Each subunit has a molecular mass of 23 to 31 kDa

and spans the membrane with sixα-helices joined

FIGURE 1.10(A) Plant aquaporins are found in the plasma membrane (PIPs) as well

as the tonoplast membrane (TIPs) of the vacuole to regulate cellular water flow.

(B) Each of the four subunits of a plant aquaporin spans the membrane six times. Both

the N-terminus (N) and C-terminus (C) are located in the cytoplasm. (C) Aquaporins

enhance water flow (thick arrow) and because they are gated allow the cell to regulate cellular flow
compared to water flow through pure lipid bilayers (thin arrow).

The interior of the aquaporin channel contains hydrophilic amino acids, which interact with water,
whereas the exterior of the protein channel consists of hydrophobic

amino acids, which interact with the lipid fatty acids of the membrane bilayer.

5 intervening loops (Figure 1.10B) in contrast to the

β-pleated sheet structure of porins. These protein subunits fold within the membrane such that the
hydrophobic amino acids are on the outer side of the pore and

interact with the hydrophobic fatty acids of the lipid

bilayer whereas the hydrophilic amino acids are in the

inner side of the pore and interact with water molecules

as they move through the pore from one side of the

membrane to the other. Aquaporins can be open or

closed to regulate the movement of water across the

membrane.Gatingis the term used to describe this regulated opening and closing of these protein
channels.

Gating through PIPs can be controlled by cytoplasmic

pH, the concentration of divalent cations such as Ca

2+

as well as by aquaporin protein phosphorylation.

As discussed above, lipid bilayers are quite permeable to water. Thus, there are two possible
pathways

for the movement of water across a membrane—one

pathway through the lipid bilayer itself and the second

pathway through an aquaporin (Figure 1.10C). If this

is so, why does a cell require any aquaporins at all?

The results of recent experiments clearly indicate that

the rate of water movement through a lipid bilayer with

aquaporins is faster than a membrane that contains lipids

only. Thus, the presence of aquaporins provides a low

resistance pathway for the movement of water across

a membrane. Furthermore, since aquaporins are gated,

this provides greater control for the movement of water

intracellularly as well as intercellularly. For example,

the water permeability of the tonoplast is two orders of

magnitude (102

times) greater than that of the plasma

membrane. Thus, this allows the vacuole to replenish or

buffer the cytoplasm with water when the cell is exposed

to hypertonic conditions. Thus, aquaporins are important in regulating the osmotic properties of
plant cells.

This process is calledosmoregulation.