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Arabidopsis LEAFY COTYLEDON2\n(LEC2) gene is a central embryonic regulator that serves\ncritical roles both early and late during embryo development.\nLEC2 is required for the maintenance of suspensor\nmorphology, specification of cotyledon identity, progression through\nthe maturation phase, and suppression of premature germination. We\ncloned the LEC2 gene on the basis of its chromosomal\nposition and showed that the predicted polypeptide contains a B3\ndomain, a DNA-binding motif unique to plants that is characteristic of\nseveral transcription factors. We showed that LEC2 RNA\naccumulates primarily during seed development, consistent with our\nfinding that LEC2 shares greatest similarity with the B3 domain\ntranscription factors that act primarily in developing seeds,\nVIVIPAROUS1/ABA INSENSITIVE3 and FUSCA3. Ectopic, postembryonic\nexpression of LEC2 in transgenic plants induces the\nformation of somatic embryos and other organ-like structures and often\nconfers embryonic characteristics to seedlings. Together, these results\nsuggest that LEC2 is a transcriptional regulator that establishes a\ncellular environment sufficient to initiate embryo development.\nEmbryogenesis in flowering\nplants begins with the double fertilization event in which the zygote\nand endosperm are formed after fusion of sperm cells with the egg cell\nand central cell of the female gametophyte, respectively. The endosperm\ninitially undergoes syncytial development with formation of\nnuclear-cytoplasmic domains, but later cellularizes (1). The developing\nembryo is nourished by the endosperm and, in many plants, only the\nperipheral layer of the endosperm remains in the mature seed.\nDevelopment of the zygote into the mature embryo can be divided\nconceptually into two distinct phases. During the early morphogenesis\nphase, the basic body plan of the plant is established with expression\nof polarity as a shoot–root axis, specification of morphological\ndomains within the embryo, and formation of embryonic tissue and organ\nsystems (2–4). The morphogenesis phase is followed by a period of\nmaturation in which processes critical for seed formation occur (5, 6).\nDuring this late phase, reserves such as storage proteins and lipids\nare synthesized at high rates and accumulate in the seed. It is also\nduring the maturation phase that the embryo acquires the ability to\nwithstand desiccation at the final stage of seed development. At the\nend of embryogenesis, the seed consists of a mature, desiccated embryo\nthat is quiescent metabolically. Although many aspects of embryogenesis\nhave been characterized extensively, little is known at a mechanistic\nlevel of the processes that initiate embryo development.\nThe Arabidopsis LEAFY COTYLEDON (LEC) genes,\nLEC1, LEC2, and FUSCA3 (FUS3), play\nkey roles in controlling embryo development (7). Unlike most other\nembryonic regulators that function during specific stages of\nembryogenesis (8–11), LEC genes are unique in that they are\nrequired for normal development during both the morphogenesis and\nmaturation phases. Early in embryogenesis, LEC genes are\nrequired to specify suspensor cell fate and cotyledon identity\n(12–16). Late in embryogenesis, LEC genes are needed during\nthe maturation phase for the acquisition of desiccation tolerance and\nthe expression of many maturation-specific genes (13–17). Consistent\nwith the finding that conditions that promote maturation suppress\ngermination (6), lec mutant embryos prematurely activate the\npostgermination program (13, 15, 16, 18). Thus, LEC genes\nplay a central role in controlling many aspects of embryogenesis, and\nthey are candidates as regulators that coordinate the morphogenesis and\nmaturation phases.\nIdentification and analysis of two LEC genes confirmed their\nregulatory role in embryogenesis and provided insight into their\nfunctions. LEC1 shares extensive sequence similarity with the HAP3\nsubunit of CCAAT-binding transcription factor, implicating LEC1 as a\ntranscriptional regulator (12). Ectopic expression of LEC1\nconfers embryonic characteristics to seedlings and results in the\nformation of embryo-like structures on the surfaces of leaves,\nsuggesting that the gene plays a role in conferring embryogenic\ncompetence to cells (12). Thus, we hypothesized that LEC1 establishes a\ncellular environment that promotes embryo development and that this\nenvironment coordinates the morphogenesis and maturation phases.\nFUS3 also encodes a regulatory protein: a B3 domain\ntranscription factor that accumulates primarily during seed development\n(19). Transient assays showed that FUS3 is sufficient to activate genes\nusually expressed during maturation (20). Thus, two LEC\ngenes seem to be involved in controlling embryo development by\nregulating transcription of other genes.\nIn this article, we focus on the LEC2 gene to determine its\nrole in embryo development. Because genetic studies suggest that\nLEC1 and LEC2 may have partially redundant\nfunctions (12, 15), it is possible that LEC2 also functions in the\ninitiation and coordination of embryo development. We cloned the\nLEC2 gene and showed that it is expressed preferentially\nduring embryogenesis and encodes a protein with similarity to other\nseed-specific transcription factors. Significant insight into the role\nof the gene was obtained by showing that transgenic plants expressing\nthe LEC2 gene ectopically form somatic embryos. Together,\nthese results indicate that LEC2 is sufficient to induce\nembryogenic competence.\n"}