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The generation of two non-identical membrane compartments via exchange ofvesicles is considered to require two types of vesicles specified by distinctcytosolic coats that selectively recruit cargo and two membrane-bound SNAREpairs that specify fusion and differ in their affinities for each type ofvesicles. The mammalian Golgi complex is composed of 6-8 non-identicalcisternae that undergo gradual maturation and replacement yet features only twoSNARE pairs. We present a model that explains how the distinct composition ofGolgi cisternae can be generated with two and even a single SNARE pair and onevesicle coat. A decay of active SNARE concentration in aging cisternae providesthe seed for a cis > trans SNARE gradient that generates the predominantlyretrograde vesicle flux which further enhances the gradient. This flux in turnyields the observed inhomogeneous steady-state distribution of Golgi enzymes,which compete with each other and with the SNAREs for incorporation intotransport vesicles. We show analytically that the steady state SNAREconcentration decays exponentially with the cisterna number. Numericalsolutions of rate equations reproduce the experimentally observed SNAREgradients, overlapping enzyme peaks in cis, medial and trans and the reportedchange in vesicle nature across Golgi: Vesicles originating from youngercisternae mostly contain Golgi enzymes and SNAREs enriched in these cisternaeand extensively recycle through the Endoplasmic Reticulum (ER), while the othersubpopulation of vesicles contains Golgi proteins prevalent in older cisternaeand hardly reaches the ER.

A model for the self-organization of vesicular flux and protein distributions in the Golgi apparatus