Sather D Noah, Golenberg Edward M
Department of Biological Sciences, Wayne State University, Detroit, MI, 48202, USA.
Planta. 2009 Feb;229(3):507-21. doi: 10.1007/s00425-008-0851-9. Epub 2008 Nov 13.
The AP1/FUL clade of MADS box genes have undergone multiple duplication events among angiosperm species. While initially identified as having floral meristem identity and floral organ identity function in Arabidopsis, the role of AP1 homologs does not appear to be universally conserved even among eudicots. In comparison, the role of FRUITFULL has not been extensively explored in non-model species. We report on the isolation of three AP1/FUL genes from cultivated spinach, Spinacia oleracea L. Two genes, designated SpAPETALA1-1 (SpAP1-1) and SpAPETALA1-2 (SpAP1-2), cluster as paralogous genes within the Caryophyllales AP1 clade. They are highly differentiated in the 3', carboxyl-end encoding region of the gene following the third amphipathic alpha-helix region, while still retaining some elements of a signature AP1 carboxyl motifs. In situ hybridization studies also demonstrate that the two paralogs have evolved different temporal and spatial expression patterns, and that neither gene is expressed in the developing sepal whorl, suggesting that the AP1 floral organ identity function is not conserved in spinach. The spinach FRUITFULL homolog, SpFRUITFULL (SpFUL), has retained the conserved motif and groups with Caryophyllales FRUITFULL homologs. SpFUL is expressed in leaf as well as in floral tissue, and shows strong expression late in flower development, particularly in the tapetal layer in males, and in the endothecium layer and stigma, in the females. The combined evidence of high rates of non-synonymous substitutions and differential expression patterns supports a scenario in which the AP1 homologs in the spinach AP1/FUL gene family have experienced rapid evolution following duplication.
MADS盒基因的AP1/FUL进化枝在被子植物物种中经历了多次复制事件。虽然最初在拟南芥中被鉴定为具有花分生组织身份和花器官身份功能,但即使在真双子叶植物中,AP1同源物的作用似乎也并非普遍保守。相比之下,FRUITFULL在非模式物种中的作用尚未得到广泛研究。我们报道了从栽培菠菜(Spinacia oleracea L.)中分离出三个AP1/FUL基因。两个基因,命名为SpAPETALA1-1(SpAP1-1)和SpAPETALA1-2(SpAP1-2),在石竹目AP1进化枝中作为旁系同源基因聚类。它们在基因的3'端羧基编码区域,即在第三个两亲性α-螺旋区域之后高度分化,同时仍保留了一些标志性AP1羧基基序的元素。原位杂交研究还表明,这两个旁系同源物已经进化出不同的时空表达模式,并且在发育中的萼片轮中均未表达,这表明AP1花器官身份功能在菠菜中并不保守。菠菜FRUITFULL同源物SpFRUITFULL(SpFUL)保留了保守基序,并与石竹目FRUITFULL同源物聚类。SpFUL在叶片以及花组织中表达,并且在花发育后期显示出强烈表达,特别是在雄性的绒毡层以及雌性的药室内壁层和柱头中。非同义替换率高和差异表达模式的综合证据支持了这样一种情况,即菠菜AP1/FUL基因家族中的AP1同源物在复制后经历了快速进化。