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Chapter 6 Axis specification The processes of the embryogenesis may “officially” begin at fertilization, but many of the molecular events critical for Drosophila embryogenesis actually occur during oogenesis. Before oogenesis, the oogonium divides four times, giving rise to 16 interconnected cells: 15 nurse cells + 1 oocyte procursor. bicoid gene encodes the morphogen(形态发生素)responsible for head structures A model of A-P pattern generation by the maternal effect genes Four maternal protein gradients in the early embryo: An anterior- to- posterior gradient of Bicoid protein An a-p gradient of Hunchback protein A p-a gradient of Nanos protein A p-a gradient of Caudal protein The terminal gene group In addition to A and P morphogens, there is the third set of maternal genes whose proteins generate the unsegmented extremities(端部)of a-p axis: the acron(原头区) and the telson(尾区). The transmembrane Torso protein (receptor) is evenly distributed throughout the plasma membrane. It is activated only at the two poles of the oocyte. Torso is a receptor tyrosine kinase (RTK,酪氨酸受体激酶). The end products of the RTK cascade inactivate Capicua protein activate +Bicoid Segmentation genes Cell fate commitment in Drosophila has two steps: Specification Determination The specification of cell fate in early development depending on cues provided protein gradients is flexible. The transition from specification to determination in Drosophila is mediated by the segmentation genes: Gap genes: mutants lacking several contiguous segments Pair-rule genes: mutants lacked portions of every other segment Segment polarity genes: mutants showed defects in every segment After the segmental boundaries have been established, the characteristic structures of each segment are specified. This specification is accomplished by the homeotic selector genes. Establishment of a-p axis 1) maternal effect genes, establish the anterior-posterior axis 2) gap ge
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