图4 甘蓝柱头乳突细胞拒绝“自花”花粉的自交不亲和分子过程Newbigin等 [ 70 ] , Deshaies等 [ 85 ] “自花”分泌的SCR存在于细胞壁中, 在SLG协助下穿过细胞壁到达细胞膜表面, 与处于非活性状态的SRK-SRK二聚体结合, 活化的SRK二聚体解聚, SRK胞内激酶域被磷酸化并瞬间磷酸化MLPK, 同时引起ARC1从细胞核定位到膜附近 [ 60 ] , ARC1 C端臂重复区同时与磷酸化的SRK激酶域、MLPK作用, ARC1活化, 其N端与Exo70A1结合, 在E1、E2共同协助下, Exo70A1被多个泛素分子泛素化, 至一定阈值时, ARC1 N端的U-box引导泛素化的Exo70A1至26S蛋白酶体处 [ 71 ] , 在酶体中心被降解, 不断被降解的Exo70A1间接或直接致MOD和MIP-MOD蛋白关闭, 柱头细胞表面缺水, 将不再向花粉运输水分等物质, 最终抑制自花花粉管的生长及花粉的萌发, 发生自交不亲和反应. Fig. 4 Brassica oleracea L. self-incompatible molecular process that papilla cell rejects self-pollen In the absence of SCR, SRK binds to the same haplotype SRK, forming SRK-SRK dimer; the protein ARC1 shuttles between the nucleas and cytoplasm. In the presence of self-pollen, SCR, released from the pollen coat, binds to and actives SRK, causing the dimer depolymerized, SRK then phosphorylatesP itself and in a moment phosphorylates MLPK, the two phosphorylated moleculars then both interactive with the C terminal of ARC1, ARM, made ARC1 phosphorylated and activated, changing location from nucleus to membrane [ 60 ] , at the same time, ARC1 N terminal attaches to Exo70A1, by the means of Ub-activation enzymes, E1, and Ub-conjugating enzymes, E2, Exo70A1 be ubiquitined by several Ubs, then ARC1 U-box leads this Exo70A1 to 26S proteasome [ 71 ] and be degraded, as the lever of Exo70A1 protein lower some threshold value causes the MOD and MIP-MOD closing, preventing water and other material from being delivered to the pollen, inhibiting the development of pollen tube and the germination of self pollen, that is the essential display of SI reaction. |