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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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SIGNALING IN PLANTS

883

(A)

(B) ABSENCE OF AUXIN

CH 2 COOH

auxin-response factor (ARF)

N

H

transcriptional repressor (Aux/IAA)

AUXIN TARGET GENES OFF

(C) PRESENCE OF AUXIN

Figure 15–71 The auxin signaling

pathway. (A) The structure of the auxin

indole-3-acetic acid. (B) In the absence of

auxin, a transcriptional repressor protein

(called Aux/IAA) binds and suppresses

a transcription regulatory protein (called

auxin-response factor, ARF), which is

required for the transcription of auxinresponsive

genes. (C) The auxin receptor

proteins are mainly located in the

nucleus and form part of ubiquitin ligase

complexes (not shown). When activated

by auxin binding, the receptor–auxin

complexes recruit the ubiquitin ligase

complexes, which ubiquitylate the Aux/IAA

proteins, marking them for degradation in

proteasomes. ARF is now free to activate

the transcription of auxin-responsive genes.

There are many ARFs, Aux/IAA proteins,

and auxin receptors that work as illustrated.

auxin

active

ARF

Aux/IAA

auxin receptor protein

ubiquitin ligase

complex

ubiquitylation

and degradation of

Aux/IAA protein

TRANSCRIPTION OF AUXIN TARGET GENES

cells via the circulatory system, auxin has its own transport system. Specific plasma-membrane-bound

influx transporter proteins and efflux transporter proteins

move auxin into and out of plant cells, respectively. The efflux transporters can be

distributed asymmetrically in the plasma membrane to make the efflux of auxin

directional. A row of cells with their auxin efflux transporters confined to the basal

plasma membrane, for example, will transport auxin from the top of the plant to

the bottom.

In some regions of the plant, the localization of the auxin transporters, and

therefore the direction of auxin flow, is highly MBoC6 dynamic m15.86/15.69

and regulated. A cell can

rapidly redistribute transporters by controlling the traffic of vesicles containing

them. The auxin efflux transporters, for example, normally recycle between intracellular

vesicles and the plasma membrane. A cell can redistribute these transporters

on its surface by inhibiting their endocytosis in one domain of the plasma

membrane, causing the transporters to accumulate there. One example occurs in

the root, where gravity influences the direction of growth. The auxin efflux transporters

are normally distributed symmetrically in the cap cells of the root. Within

minutes of a change in the direction of the gravity vector, however, the efflux transporters

redistribute to one side of the cells, so that auxin is pumped out toward the

side of the root pointing downward. Because auxin inhibits root-cell elongation,

this redirection of auxin transport causes the root tip to reorient, so that it grows

downward again (Figure 15–72).

Phytochromes Detect Red Light, and Cryptochromes Detect Blue

Light

Plant development is greatly influenced by environmental conditions. Unlike

animals, plants cannot move when conditions become unfavorable; they have

to adapt or they die. The most important environmental influence on plants is

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