Tapetal development and abiotic stress: a centre of vulnerability

Many self-fertilising crops are particularly sensitive to abiotic stress at the reproductive stage. In rice (Oryza sativa L.) and wheat (Triticum aestivum L.), for example, abiotic stress during meiosis and the young microspore stage indicates the tapetum is highly vulnerable and that the developmental program appears to be compromised. Tapetal hypertrophy can occur as a consequence of cold and drought stress, and programmed cell death (PCD) is delayed or inhibited. Since the correct timing of tapetal PCD is essential for pollen reproduction, substantial losses in grain yield occur. In wheat and rice, a decrease in tapetal cell wall invertase levels is correlated with pollen abortion and results in the amount of hexose sugars reaching the tapetum, and subsequently the developing microspores, being severely reduced ('starvation hypothesis'). ABA and gibberellin levels may be modified by cold and drought, influencing levels of cell wall invertase(s) and the tapetal developmental program, respectively. Many genes regulating tapetal and microspore development have been identified in Arabidopsis thaliana (L.) Heynh. and rice and the specific effects of abiotic stresses on the program and pathways can now begin to be assessed.