Chemistry and biology of boron.

Boron is an essential nutrient for certain organisms, notably vascular plants and diatoms. Cyanobacteria require boron for formation of nitrogen-fixing heterocysts and boron may be beneficial to animals. Boron deficiency in plants produces manifold symptoms: many functions have been postulated. Deficiency symptoms first appear at growing points, within hours in root tips and within minutes or seconds in pollen tube tips, and are characterized by cell wall abnormalities. Boron-deficient tissues are brittle or fragile, while plants grown on high boron levels may have unusually flexible or resilient tissues. Borate forms cyclic diesters with appropriate diols or polyols. The most stable are formed with cis-diols on a furanoid ring. Two compounds have this structure physiologically: ribose in ribonucleotides and RNA, and apiose in the plant cell wall. Germanium can substitute for boron in carrot cell cultures. Both boron and germanium are localized primarily in the cell wall. We postulate that borate-apiofuranose ester cross-links are the auxin-sensitive acid-growth link in vascular plants, that the cyanobacterial heterocyst envelope depends on borate cross-linking of mannopyranose and/or galactopyranose residues in a polysaccharide-lipid environment, and that boron in diatoms forms ester cross-links in the polysaccharide cell wall matrix rather than boron-silicon interactions. Complexing of ribonucleotides is probably a factor in boron toxicity.