非洲狼尾草无融合生殖胚胎学研究

EMBRYOLOGICAL STUDIES ON APOMIXIS IN PENNISETUM SQUAMULATUM

报道非洲狼尾草（ＰｅｎｎｉｓｅｔｕｍｓｑｕａｍｕｌａｔｕｍＦｒｅｓｅｎ）的胚囊形成、胚胎发生与发育过程。非洲狼尾草的孢原细胞直接发育成大孢子母细胞，并由它分裂产生三分体。从大孢子母细胞发育至三分体的不同阶段，均会出现败育。性细胞退化期间，其周围的珠心组织中，常出现一至多个体积较大的无孢子生殖原始细胞。通常只有靠近珠孔端的１个无孢子生殖原始细胞体积进一步增大，并出现大液泡，发育成无孢子生殖单核胚囊。随后，其核经连续两次有丝分裂，形成无孢子生殖四核胚囊，胚囊内的４个核常聚积在珠孔端，４个核进一步分化形成１个卵细胞、１个助细胞和具两个极核的中央细胞，没有反足细胞。胚囊发育属于大黍型。其它的无孢子生殖原始细胞能发育到单核或二核胚囊阶段，而后核解体导致胚囊败育。胚的发生有两种类型：（１）早发生胚。大多数胚囊在开花前一、二天，次生核未分裂，卵细胞不经受精，自发分裂形成胚。（２）迟发生胚。少数胚囊的卵细胞不经过受精，但需要在开花后三、四天次生核分裂为多个胚乳核时才开始分裂。无论是早发生胚或迟发生胚，卵细胞在分裂前具有极性，珠孔端有大液泡，细胞质稀薄，合点端细胞质较浓。胚的发育经历球形胚、梨形胚和胚分化阶段。

Studies on the formation and development of the embryo sac of the apomictic material of Pennisetum squamulatum Fresen indicated that normal archesporial cell did form with consequent development of a megaspore mother cell and later meiotic division to give rise to a triad. But invariably the megaspore mother cell and the triad underwent degeneration after formation. During the period of formation or degeneration of the megaspore or the triad a number of nucellar cells around the degenerated sexual cell became much enlarged. Frequently, one of the enlarging nucellar cells near the micropylar end became vacuolated and then developed into an aposporous uninucleate embryo sac, which underwent two further mitotic divisions to form an aposporous fournucleate embryo sac, where the four nuclei remained in the micropylar end. Thus in the mature aposporous embryo sac there were one egg cell, one synergid and one central cell (containing two polar nuclei). Antipodal cells were completely lacking. The pattern of development of the aposporous embryo sac resembles the panicum type. There were two types of embryo formed during apomictic development namely (1) The pregenesis embryo—embryo fomed without fertilization,1 to 2 days before anthesis, and (2) The lategenesis embryo—derived from the unfertilized egg cells, 3 to 4 days after anthesis. In the lategenesis embryo type, the egg cell divided after the secondary nucleus has undergone division to form the endosperm nuclei. All egg cells developed vacuoles before they differentiated into embryos. The development of the aposporous embryo followed the sequence of the formation of globular, pearshaped embryo and full stages of differentiation. The unfertilized secondary nucleus divides to form free endosperm nuclei after being stimulated by pollination. The development of the endosperm belongs to the nucleartype.