Tansley Review No. 108 Molecular events of vesicle trafficking and control by SNARE proteins in plants

Abstract

CONTENTS

Summary 389

I. INTRODUCTION 389

II. 1. The SNARE hypothesis 393

2. SNAREs and associates 395

(a) NSF 395

(b) SNAP 395

(c) SNARE 395

(d) Rab and Rab-associated proteins 397

(e) Other interacting proteins 398

(f) Clostridial neurotoxins 399

3. New perspectives on SNARE function 400

(a) Revisiting the SNARE hypothesis 400

(b) Roles for SNARE proteins 401

III. VESICLE TRAFFICKING IN PLANTS 402

1. Identifying secretory proteins in plants 402

2. Placing syntaxins in order 403

3. Functional analysis of plant SNAREs in vesicle trafficking 404

4. Small GTPases 405

5. Searching for a vesicle–fusion complex 405

IV. REGULATION OF VESICLE TRAFFICKING IN PLANT CELLS 406

1. Hormonal control of secretion 406

(a) Auxin-evoked secretion and cell growth 406

(b) Gibberellic acid and secretion by barley aleurone 407

(c) Abscisic acid control of stomatal aperture 407

2. Coupling secretion to cytosolic-free Ca⁺ concentration 408

3. Stress-evoked secretion 409

(a) Mechanical stress 409

(b) Wounding and pathogen attack 409

V. CONCLUSION 410

Acknowledgements 411

References 411

Eukaryotic cells share a set of secretory pathways for the flux of membrane and protein material. In 1993, ideas about the functioning of three major proteins of the neurosecretory complex were consolidated in the SNARE hypothesis, which proposed that the interaction of these proteins provides both the specificity for vesicle targeting and the molecular machinery for fusion between vesicle and target membranes. Subsequetly, the organization, molecular mechanics and control of vesicle trafficking have become topics of intense research, and the hypothesis has evolved to accommodate new discoveries from the analysis of secretion in yeast and mammals. It is likely to be challenged again as more information comes to light about secretory processes in plants. New tools for measuring and manipulating vesicle traffic and secretion are now being used, drawing on in vivo fluorescence and capacitance recording as well as genetic engineering. These new technologies have already begun to yield details wholly unexpected from past studies. Here we focus on recent findings relating to the mechanisms of vesicle trafficking and the background to these developments, highlighting both current understanding of the molecular events of secretion and the gaps therein, as well as discussing emerging themes from work with plants.