Barley grain development toward an integrative view.

Seeds are complex structures composed of several maternal and filial tissues which undergo rapid changes during development. In this review, the barley grain is taken as a cereal seed model. Following a brief description of the developing grain, recent progress in grain development modeling is described. 3-D/4-D models based on histological sections or nondestructive NMR measurements can be used to integrate a variety of datasets. Extensive transcriptome data are taken as a frame to augment our understanding of various molecular-physiological processes. Discussed are maternal influences on grain development and the role of different tissues (pericarp, nucellus, nucellar projection, endosperm, endosperm transfer cells). Programmed cell death (PCD) is taken to pinpoint tissue specificities and the importance of remobilization processes for grain development. Transcriptome data have also been used to derive transcriptional networks underlying differentiation and maturation in endosperm and embryo. They suggest that the "maturation hormone" ABA is important also in early grain development. Massive storage product synthesis during maturation is dependent on sufficient energy, which can only be provided by specific metabolic adaptations due to severe oxygen deficiencies within the seed. To integrate the great variety of data from different research areas in complex, predictive computational modeling as part of a systems biology approach is an important challenge of the future. First attempts of modeling barley grain metabolism are summarized.