Pollenkitt is secreted by the tapetum of the anther
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Peer-reviewed literature establishes that the pollen coat, or pollenkitt, is largely derived and secreted from the tapetum cells within the anther during pollen development.
<h4>Background</h4>The male gametophyte developmental programme can be divided into five phases which differ in relation to the environment and pollen hydration state: (1) pollen develops inside the anther immersed in locular fluid, which conveys substances from the mother plant--the microsporogenesis phase; (2) locular fluid disappears by reabsorption and/or evaporation before the anther opens and the maturing pollen grains undergo dehydration--the dehydration phase; (3) the anther opens and pollen may be dispersed immediately, or be held by, for example, pollenkitt (as occurs in almost all entomophilous species) for later dispersion--the presentation phase; (4) pollen is dispersed by different agents, remaining exposed to the environment for different periods--the dispersal phase; and (5) pollen lands on a stigma and, in the case of a compatible stigma and suitable conditions, undergoes rehydration and starts germination--the pollen-stigma interaction phase.<h4>Scope</h4>This review highlights the issue of pollen water status and indicates the various mechanisms used by pollen grains during their five developmental phases to adjust to changes in water content and maintain internal stability.<h4>Conclusions</h4>Pollen water status is co-ordinated through structural, physiological and molecular mechanisms. The structural components participating in regulation of the pollen water level, during both dehydration and rehydration, include the exine (the outer wall of the pollen grain) and the vacuole. Recent data suggest the involvement of water channels in pollen water transport and the existence of several molecular mechanisms for pollen osmoregulation and to protect cellular components (proteins and membranes) under water stress. It is suggested that pollen grains will use these mechanisms, which have a developmental role, to cope with environmental stress conditions.
I have used cellular and molecular genetic and bioinformatic approaches to characterise the components of the pollen coat in plants of the family Brassicaceae, including Arabidopsis thaliana and several brassicas including Brassica napus, B. oleracea, and B. rapa. The pollen coat in these species is mostly made up of a unique mixture of lipids that is highly enriched in acylated compounds, such as sterol esters and phospholipids. These acyl lipids are characterised by an unusually high degree of saturation. The fatty acids typically contain 70-90% saturated acyl residues such as myristate, palmitate, and stearate. The major sterol components of the pollen coat are saturated fatty acyl esters of stigmasterol, campesterol, and campestdienol. In addition to lipids, the second major component of the pollen coat is a specific group of proteins that is dominated by a family of proteins that we term pollenins. Although pollenins are by far the major protein components of the pollen coat of members of the Brassicaceae, proteomic analysis reveals that there are several additional protein components, including lipases, protein kinases, a pectin esterase, and a caleosin. The biosynthesis of these lipids and proteins and their significance for overall pollen function are reviewed and discussed.
The tapetum is a single layer of secretory cells which encloses the anther locule and sustains pollen development and maturation. Upon apoptosis, the remnants of the tapetal cells, consisting mostly of lipids and proteins, fill the pits of the sculpted exine to form the bulk of the pollen coat. This extracellular matrix forms an impermeable barrier that protects the male gametophyte from water loss and UV light. It also aids pollen adhesion and hydration and retains small signaling compounds involved in pollen-stigma communication. In this study, we have updated the list of the pollen coat's protein components and also discussed their functions in the context of sexual reproduction.
The pollen coat is the outermost domain of the pollen grain and is largely derived from the anther tapetum, which is a secretory tissue that degenerates late in pollen development. By being localised at the interface of the pollen-stigma interaction, the pollen coat plays a central role in mediating early pollination events, including molecular recognition. Amongst species of the Brassicaceae, a growing body of data has revealed that the pollen coat carries a range of proteins, with a number of small cysteine-rich proteins (CRPs) being identified as important regulators of the pollen-stigma interaction. By utilising a state-of-the-art liquid chromatography/tandem mass spectrometry (LC-MS/MS) approach, rich pollen coat proteomic profiles were obtained for <i>Arabidopsis thaliana</i>, <i>Arabidopsis lyrata</i>, and <i>Brassica oleracea</i>, which greatly extended previous datasets. All three proteomes revealed a strikingly large number of small CRPs that were not previously reported as pollen coat components. The profiling also uncovered a wide range of other protein families, many of which were enriched in the pollen coat proteomes and had functions associated with signal transduction, cell walls, lipid metabolism and defence. These proteomes provide an excellent source of molecular targets for future investigations into the pollen-stigma interaction and its potential evolutionary links to plant-pathogen interactions.
This special mini-review was planned as a synthesis of current understanding of the role of tapetum and orbicules, of the knowledge on pollenkitt, with addition of our own data on experimental orbicule simulation. The aim was to show the development of knowledge and ideas through time. Tapetum types are so changeable that the idea of norm becomes ghostly. The review is based on our own and other authors' results. Cyclic-invasive tapeta, surprising exine-like tapetal surface, direct connections of tapetum with microspores via filaments are probably not rare phenomena. Our in vitro experiments on microspore exine simulations, which have led also to appearance of orbicule-like structures, support the view of their by-product nature, based on self-assembly. Different types of orbicules and their development are examined. Tapetum and orbicule functions and especially pollenkitt production are reviewed, together with the data on sporopollenin. Some concise data on molecular and genetic studies are added.
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