Collaborative Research: Hybrid Seed Inviability and the Evolution of Endosperm Development in Mimulus

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The seed of flowering plants contains two very important structures: the embryo that will become a seedling and eventually grow into the mature plant, and the endosperm, a tissue that provides nutrients to the embryo as it grows. If you have eaten rice, you have eaten endosperm, the starchy portion that fills the majority of the rice grain. Since the emergence of agriculture, humans have selected and bred seeds with a larger and starchier endosperm portion, ultimately creating the crop varieties we now depend upon. Endosperm, either directly or indirectly through animal feed, now provides an estimated 67% of the calories consumed by humans worldwide, and improving on these varieties remains of fundamental importance. Thus, from both an agricultural and food security perspective the development of the endosperm is critical to humankind's survival. This research will investigate the genes that control endosperm development and the molecular and evolutionary interactions between the endosperm and the embryo in the monkey flower (genus Mimulus). Although not eaten by humans, due to its ecological and evolutionary history, Mimulus provides a unique opportunity to study evolutionary and molecular aspects of the interactions between endosperm and embryo. As these interactions regulate the growth of the endosperm and thus the size of the nutritious portion of the seed in agricultural varieties, the results of this research in Mimulus are expected to engender a molecular understanding of a developmental process critical to human agricultural productivity. This research will inspire new high school biology teaching modules in genetics, evolution, and plant developmental biology that highlight the importance of the endosperm to humankind's diet, as well as the roles of pollinators and reproductive barriers in generating the diversity of life through the evolutionary process.Recent evidence links the evolutionary theory of parent-offspring conflict over maternal resources to the evolution of epigenetically imprinted genes controlling normal seed endosperm development. In interploidy hybrids, dosage-sensitivity of paternal and maternal allelic expression in developing endosperm is critical to normal development, and disruption leads to seed abortion. As yet it remains unknown whether such conflict over embryo provisioning leads to evolutionary divergence of imprinted genes or their targets, and ultimately hybrid seed lethality, in diploid species. The investigators propose to test this possibility by determining the genetic basis and evolutionary mechanisms underlying species divergence in endosperm development and leading to hybrid seed abortion, a principal isolating barrier in the Mimulus guttatus sp. complex. This research integrates genomic, molecular genetic and developmental biology approaches, utilizing high-throughput genome mapping, RNA sequencing technologies, and transgenic techniques to elucidate the genetic mechanisms contributing to seed abortion in M. guttatus x M. nudatus crosses. In so doing, it examines questions of fundamental interest to evolutionary biologists (how are new species formed?), and to developmental biologists (how is endosperm growth regulated?) while addressing a question of interest to both: what molecular changes might rapidly evolve when the mother's need to regulate growth and development differs from that of her offspring? Answering these questions will shed light on the developmental processes regulating endosperm growth and the evolutionary mechanisms driving divergence in flowering plants.