Sokolov D D, Timonin A K
Zh Obshch Biol. 2007 Mar-Apr;68(2):83-97.
Molecular phylogenetic data have drastically changed the views on the phylogeny of higher plants. All the extant gymnosperms were asserted as a monophyletic group opposed to the highly isolated angiosperms. The 'Anthophyte Theory' was thus rejected. The identification and analysis of gymnosperm orthologues of genes regulating flower development in angiosperms resulted in the formulation of the 'Mostly Male Theory' of the evolutionary origin of flower; this theory does not contradict the concept of monophyly of all the extant gymnosperms. The Mostly Male Theory assumes that the origin of angiosperms was caused by a loss of the Needle family gene that effected ovuliferous (female) organs and the translocation of the ovules onto the adaxial side of some of the (male) leafy microsporangiophores. Having acquired ovules, the former microsporangiophores started evolving into the carpels. The prerequisite bisexual design of the ancestral fructification thus becomes unnecessary. Indeed, this assumption suggests the deriving of Angiosperms from any gymnosperm plant with leafy microsporangiophores. The problem of carpel origin has subsequently changed to some degree into the problem of the origin of the bitegmic anatropous ovule presumably inherent in ancestral Angiosperms. The Mostly Male Theory consideredeither Corystospermataceae (= Umkomasiaceae) or Caytoniaceae to be the forerunners of such an ovule. Yet the capsules of Corystospermataceae distinctly differ from angiosperm ovules in the locations of their adaxial/abaxial sides, while Caytoniaceae had no leafy microsporangiophores. This inconsistency suggests that functions of the Needle family regulatory genes in Gymnosperms should be much better understood to appraise properly both the possibilities and the consequences of their hypothetical loss by the emerging angiosperms. Moreover, the extant gymnosperm groups are actually held as monophyletic and contrasted to Angiosperms on the basis of analysing the unrepresentative scant remnants of these, mostly extinct, taxa. Therefore, traditional botanical and paleobotanical data should not be rejected. In any case, Meyen's idea angiosperms origin from Bennettitales is worth being retained as a hypothesis to be tested with new results of both paleobotany and molecular biology.
分子系统发育数据极大地改变了人们对高等植物系统发育的看法。所有现存的裸子植物被认定为一个单系类群，与高度孤立的被子植物相对。因此，“花植物理论”被摒弃。对调控被子植物花发育的基因的裸子植物直系同源基因进行鉴定和分析，得出了花进化起源的“大多为雄性理论”；该理论与所有现存裸子植物单系性的概念并不矛盾。大多为雄性理论认为，被子植物的起源是由于影响具胚珠（雌性）器官的针叶家族基因的缺失，以及胚珠转移到一些（雄性）具叶小孢子囊托的近轴面。获得胚珠后，原来的小孢子囊托开始进化成心皮。因此，祖先果实的双性设计前提变得不再必要。实际上，这一假设表明被子植物起源于任何具有具叶小孢子囊托的裸子植物。心皮起源问题随后在一定程度上转变为可能存在于被子植物祖先中的双珠被倒生胚珠的起源问题。大多为雄性理论认为科里斯托精子科（=乌姆科马西亚科）或凯托尼亚科是这种胚珠的前身。然而，科里斯托精子科的蒴果在其近轴/远轴面位置与被子植物胚珠明显不同，而凯托尼亚科没有具叶小孢子囊托。这种不一致表明，为了正确评估新出现的被子植物假设性缺失这些基因的可能性和后果，需要更好地理解裸子植物中针叶家族调控基因的功能。此外，基于对这些大多已灭绝类群的非代表性少量残余进行分析，现存的裸子植物类群实际上被视为单系类群，并与被子植物形成对比。因此，传统植物学和古植物学数据不应被摒弃。无论如何，迈恩关于被子植物起源于本内苏铁目的观点值得作为一个有待古植物学和分子生物学新结果检验的假设保留下来。
