葉綠體DNA
菸草的葉綠體DNA基因組，標示於內側的基因位於DNA的同一股上，標示於外側者則位於另一股，基因中的缺口為內含子。
葉綠體DNA（Chloroplast DNA，cpDNA）為真核生物細胞中葉綠體內的DNA。葉綠體等质粒体中具有一獨立於細胞核的基因組，且含有核糖體可轉譯合成自身的蛋白質[1][2]，為一半自主的胞器，葉綠體DNA最早於1959年透過生化實驗發現[3]，並於1962年透過電子顯微鏡實際觀測確認[4]。1986年菸草與地錢的葉綠體DNA被定序發表，為最早完成定序的葉綠體基因組[5][6]，目前已有大量陆生植物與藻類的质粒体被定序發表。
基因组結構
[编辑]葉綠體DNA為環狀，長度一般介於120至170kb之間[7][8][9]，分子量為8000萬至1.3億Da[10]，多數植物的葉綠體包含約120個基因[11][12]，大多編碼光合作用所需的蛋白（包括RuBisCO的大次單元）與基因表現所需的蛋白[13]，如4種rRNA、約30種tRNA、21種核糖體蛋白以及葉綠體RNA聚合酶的4個次單元[13]。絕大多數生物的葉綠體DNA均不分段，雙鞭毛蟲門的藻類則為罕見例外，此類生物的葉綠體DNA包括約40段質體，每個質體長2至10kb，包含1至3個基因，也有些質體不編碼任何基因[14]。
多數葉綠體DNA包括兩段反向重複序列（IRa與IRb），將葉綠體DNA分成大單拷貝區（LSC）與小單拷貝區（SSC）兩區域[9]。反向重複序列長4至25kb，其中植物的一般在20至25kb之間[15]，IRa與IRb的序列一般會因協同演化而非常近似[14]。植物葉綠體DNA的反向重複序列演化上相當保守[9][15]，藍菌基因組與灰藻和紅藻的葉綠體基因組中也有與其同源的序列，顯示此序列在演化上的起源很早，在葉綠體出現前即已存在[14]。有些葉綠體（豌豆與數種紅藻[14]）的基因組丟失了反向重複序列[15][16]，紫菜屬紅藻葉綠體則有一個重複序列發生了倒轉，使IRa與IRb變為同向排列[14]。反向重複序列可能可幫助維持葉綠體DNA的穩定[16]。
陸生植物新葉的葉綠體中一般有約100個葉綠體DNA，老葉中則僅剩15至20個[17]，多包裹成擬核，一個葉綠體中常有數個擬核[10]。葉綠體DNA雖不與組蛋白結合[18]，但紅藻的葉綠體DNA可編碼和組蛋白相似的組蛋白樣葉綠體蛋白（HC）與自身結合[19]。較原始的紅藻Cyanidioschyzon merolae（屬溫泉紅藻綱）的葉綠體擬核集中在葉綠體基質的中央，綠藻與陆生植物的葉綠體擬核則均勻分布於基質中[19]。
參見
[编辑]參考文獻
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