蘭科植物的花被由三片萼瓣、兩片側花瓣和中央花瓣特化的唇瓣所組成。研究報導顯示唇瓣對蘭花授粉扮演很重要的角色，蘭科植物的演化與其具有兩側對稱性的花朵有密切相關，而唇瓣的發育更是造成蘭科植物由擬蘭亞科具似百合輻射花形演化成兩側對稱花形的主要原因之ㄧ。本論文探討蘭花B群及E群MADS box基因參與蘭花花器決定的功能，並提出同源性蘭花花被模式 (Homeotic Orchid Tepal, HOT model) 進一步闡釋它們在蘭花花器發育所扮演之決定功能。首先，廣泛地研究分佈於5個亞科12個不同屬的蘭花B群基因。發現蘭科植物皆具有由二次基因複製產生的AP3A1/A2和AP3B1/B2四個分群以及PI同源性基因，具特化唇瓣蘭花之AP3A2同源基因在花瓣和唇瓣有差異性表現，且具有唇瓣專一性，並利用蘭花三唇瓣突變株驗證AP3A2基因的唇瓣專一性表現。其中不具顯著形態差異唇瓣之擬蘭亞科雖保有原始AP3A2同源基因，但是其可能尚未演化調控特化唇瓣的功能。基於這些研究結果，提出調控蘭花花被的Homeotic Orchid Tepal (HOT) model，說明由演化複製所產生之B群花器決定基因可參與蘭花花被之形成，特別是特化花器─唇瓣之發育。進一步研究蝴蝶蘭的四個E群基因PeSEPs，探討其基因表現、以酵母菌雙雜合實驗測試蛋白質交互作用，並以病毒誘導基因靜默檢驗其基因功能。PeSEP基因可參與或藉由影響B群基因表現調控表皮細胞形態大小、花青素及葉綠素形成，及表皮角質蠟生成而決定蘭花花萼、花瓣和唇瓣的發育。綜合以上，本研究釐清蘭花B和E群MADS box花器決定基因的分子調控關係，並了解其參與蘭花花被形態發生的重要功能。

The typical orchid floral morphology architecturally consists of three sepals, two petals, together with the most sophisticated lip, and colum. Previous studies reveal that lip plays a critical role in orchid pollination strategies. Orchid flower, remarkable for its zygomorphic syndrome, is intently related to the Orchidaceae evolutionary history. The evolution of lip is one of the major factors for orchid flower exhibiting zygomorphy to derived from the actinomorphy lily-like flower. This thesis investigates the B- and E-class MADS genes in determining orchid floral organ identities, and the Homeotic Orchid Tepal (HOT) model is proposed for explicating floral morphogenesis in Orchidaceae. Tweenty seven AP3-like and 14 PI-like genes from 12 species of orchids were widely identified. Homologous genes of PI clade and four duplicated AP3A1, AP3A2, AP3B1, and AP3B2 subclades were conserved in the orchid family. Among the four AP3 subclades, the lip specificity of orchid AP3A2 orthologs was demonstrated by the differential transcript between petal and lip of normal flowers and by the ectopic expression in lip-like petal of orchids with peloric flowers. Although an AP3A2-like gene was identified in lily-like Neuwiedia flower without obviously modified lip, the primitive AP3A2 maybe necessary but not sufficient for regulating lip development. Based on these results, the HOT model was proposed, in which dualistic characters of duplicated B-class MADS-box genes are involved in orchid perianth development and growth, especially the formation of the modified petal, lip. Furthermore, I functionally characterized the PeSEP genes that are uniformly expressed in all floral organs in respects of protein-protein interaction and gene expression by using of yeast two-hybrid and VIGS. PeSEPs themselves and possibly via regulating their downstream B-class genes involved in stimulating cell shape, cellular ultra-structures, floral pigmentation and cutin biosynthesis, but inhibiting features of leaf-like structures to determine the development of orchid sepals, petals and lip. Together, these studies have clarified the relationship of orchid B- and E-class MADS-box genes involved in the regulation of orchid floral organ identities and uncovered the critical roles in the obscure mystery of the orchid perianth morphogenesis.

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