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   » » Wiki: Sporopollenin
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Sporopollenin is a biological found as a major component of the tough outer (exine) walls of and grains. It is chemically very stable and has been described as the "toughest material in the plant kingdom". It is well preserved in and and with it surviving in spores from the mid‐ (475 million years ago) providing the earliest evidence of on land.

The exine layer is often intricately sculptured in species-specific patterns, allowing material recovered from (for example) lake sediments to provide useful information to about past plant and fungal populations. Sporopollenin has found uses in the field of as well as a marker of past () levels in the . Sporopollenin is also found in the cell walls of several taxa of , including (an ) and .

are dispersed by many different environmental factors, such as wind, water or animals. In suitable conditions, the sporopollenin-rich walls of pollen grains and spores can persist in the fossil record for hundreds of millions of years, since sporopollenin is resistant to chemical degradation by organic and inorganic chemicals.


Chemical composition
The chemical composition of sporopollenin has long been elusive due to its unusual chemical stability, in and resistance to degradation by and strong chemical reagents. It was once thought to consist of polymerised , but the application of more detailed analytical methods since the 1980s has shown that this is not correct. Analyses have revealed a complex , containing mainly long-chain fatty acids, , and traces of carotenoids in a random co-polymer. Sporopollenin likely derives from several precursors that are chemically to form a rigid structure. There is also good evidence that the chemical composition of sporopollenin is not the same in all plants, indicating it is a class of compounds rather than having one constant structure.

In 2019, thioacidolysis degradation and solid-state NMR was used to determine the molecular structure of sporopollenin, finding it primarily composed of polyvinyl alcohol units alongside other aliphatic , all crosslinked through a series of linkages. Its complex and heterogeneous chemical structure gives some protection from the biodegradative enzymes of , and animals. Some aromatic structures based on and were also identified within the sporopollenin polymer. These can absorb (UV) light, preventing it from penetrating further into the spore. This is relevant to the role of pollen and spores in transporting and dispersing of plants. The of the gametes is readily damaged by the ultraviolet component of . Sporopollenin thus provides some protection from these damages as well as a physically robust container.

Analyses of sporopollenin from the in the late 1980s have shown distinct structural differences from that of flowering plants. In 2020, more detailed analysis of sporopollenin from Lycopodium clavatum provided more structural information. It showed a complete lack of aromatic structures and the presence of a backbone of poly tetraketide-like monomers with pseudo-aromatic 2-pyrone rings. These were crosslinked to a poly(hydroxy acid) chain by ether linkages to form the polymer.


Biosynthesis
Electron microscopy shows that the that surround the developing pollen grain in the have a highly active system containing globules. These globules are believed to contain sporopollenin precursors. Tracer experiments have shown that is a major precursor, but other carbon sources also contribute. The biosynthetic pathway for phenylpropanoid is very active in tapetal cells, supporting the idea that its products are needed for sporopollenin synthesis. Chemical inhibitors of pollen development and many mutants affect the secretion of these globules by tapetal cells.


Ultraviolet protection and paleoclimatology
Sporopollenin contain that absorb both , acting to protect internal cell components including against ultraviolet damages. Plants regulate production of these phenolic components given UV exposure, making them a paleoclimatological marker for past UV levels.


See also


Further reading
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