Arabidopsis APETALA2 (AP2) encodes a member of the AP2͞EREBP (ethylene responsive element binding protein) class of transcription factors and is involved in the specification of floral organ identity, establishment of floral meristem identity, suppression of floral meristem indeterminancy, and development of the ovule and seed coat. Here, we show that loss-of-function ap2 mutations cause an increase in seed mass relative to that of wild-type seeds. Analysis of an allelic series of ap2 mutations showed that increases in seed mass corresponded with the severity of defects in flower structure, indicating that AP2 activity directly influences seed mass. Experiments with male-sterile plants and deflowered wild-type plants showed that reduced fertility of ap2 mutant plants due to abnormal flower structure accounted for only part of the increase in seed mass caused by strong ap2 mutant alleles. Reciprocal cross experiments showed that AP2 acts maternally to control seed mass. The maternal effect of AP2 on seed mass involves the regulation of both embryo cell number and cell size. We show further that ap2 mutations cause changes in the ratio of hexose to sucrose during seed development, opening the possibility that AP2 may control seed mass through its effects on sugar metabolism. Together, these results identify a role for AP2 in controlling seed mass.ap2 ͉ Arabidopsis ͉ seed size ͉ sugar metabolism S eeds of higher plants consist of three major components, each with a different genotype. The embryo that develops into the vegetative plant is diploid with a zygotic complement of genomes contributed by its parents. The endosperm, a tissue system that serves a nutritive role for the developing embryo and͞or germinating seedling, is triploid with two and one genome equivalents, respectively, contributed by the maternal and paternal parent. By contrast, the seed coat that surrounds the embryo and endosperm is strictly of maternal origin. Growth and development of the embryo, endosperm, and seed coat must be coordinated to produce the mature seed. Although seeds have been studied extensively, many aspects of seed development are not well understood, including the mechanisms that underlie seed size or mass.A critical factor in determining plant fitness is seed mass. Seed mass is negatively correlated with the number of seeds produced and positively correlated with seedling survival (1-5). Smallseeded plants are considered to be efficient colonizers because they produce large numbers of seeds, whereas seedlings of large-seeded plants are thought to more effectively withstand resource restrictions and abiotic stresses. Moreover, seed mass can vary intraspecifically in response to environmental cues, although little is known of the specific regulatory processes involved.Several factors that influence seed mass have been identified. For example, quantitative trait loci (QTL) that influence seed mass have been mapped in a number of crop plants (6)(7)(8)(9)(10)(11)(12)). An analysis of genetic factors affecting Arabidopsis seed mas...