壶瓶枣大小孢子发生与雌雄配子体发育

doi:10.16420/j.issn.0513-353x.2025-0560

摘要/Abstract

摘要：

以壶瓶枣（Ziziphus jujuba‘Hupingzao’）为试材，采用石蜡切片技术，观察大小孢子发生及雌雄配子体发育进程，为壶瓶枣花发育和育种提供理论依据。壶瓶枣花药具4室，花药壁由1层表皮、1层药室内壁、2层中层和1层分泌型绒毡层组成，发育类型为基本型；小孢子母细胞胞质分裂为同时型，经减数分裂形成四分体，呈四面体型排列；成熟花粉为2细胞型，具3个萌发孔。胚珠为倒生型；大孢子母细胞经减数分裂形成四分体，呈纵向线形排列，合点端大孢子发育为功能大孢子，雌配子体发育经历二核、四核胚囊阶段，最终形成7细胞8核的成熟胚囊，发育类型为蓼型。雌雄异熟，雄蕊发育早于雌蕊。

关键词: 枣, 壶瓶枣, 小孢子发生, 大孢子发生, 雄配子体, 雌配子体

Abstract:

Observing sporogenesis and gametogenesis is a fundamental step in revealing the mechanism of embryo abortion in Ziziphus jujuba‘Hupingzao’. The occurrence of megasporogenesis，microsporogenesis as well as the development of female and male gametophyte of‘Hupingzao’were studied by paraffin sectioning technique in this paper. Each anther has four pollen sacs. The anther wall is composed of an epidermis，an endothecium，two middle layers，and a secretory tapetum，and its development type is the basic type. Following meiosis，the microspore mother cells forme tetrahedral tetrads through simultaneous cytokinesis. The mature pollen grains are two-celled with three pollen apertures. The ovule is anatropous，and the ovary is bilocular with one ovule per locule. The megaspore mother cell undergo meiosis to form a linear tetrad along the longitudinal axis. The chalazal megaspore develops into the functional megaspore. The female gametophyte develops through binucleate and four-nucleate embryo sac stages，ultimately formed a mature 7-celled，8-nucleate embryo sac. Embryo sac development is the polygonum type.‘Hupingzao’exhibits dichogamy，with stamens developing earlier than pistils.

Key words: Ziziphus jujuba, Hupingzao, microsporogenesis, megasporogenesis, male gametophyte, female gametophyte

郭红彦, 武晋帆, 向敏, 蒋少东, 赵爱玲. 壶瓶枣大小孢子发生与雌雄配子体发育[J]. 园艺学报, 2025, 52(12): 3157-3166.

GUO Hongyan, WU Jinfan, XIANG Min, JIANG Shaodong, ZHAO Ailing. Sporogenesis and Gametogenesis of Ziziphus jujuba‘Hupingzao’[J]. Acta Horticulturae Sinica, 2025, 52(12): 3157-3166.

图/表 6

图1 壶瓶枣花药壁的发育过程 A：4个角隅处细胞分裂，箭头所指为药隔；B：箭头所指为4个花粉囊；C：箭头所指为两药室；D：初生周缘细胞和初生造孢细胞；E：表皮与药室内壁；F：完整花药壁结构；G：中层退化；H：绒毡层退化；I：药室内壁仅留有纤维层；J：纤维层破裂散出花粉粒。ppc：初生周缘细胞；psc：初生造孢细胞；ssc：次生造孢细胞；epi：表皮；end：药室内壁；ml：中层；tap：绒毡层

Fig. 1 Development process of anther wall of Ziziphus jujuba‘Hupingzao’ A：Cell division at the four corners，the arrow refers to the septum；B：The arrow refers to four pollen sacs；C：The arrow refers to two locules；D：Primary peripheral cells and primary sporogenous cells；E：Epidermis and drug chamber wall；F：Complete anther wall structure；G：Middle layer degradation；H：Tapetal degeneration；I：There is only fiber layer on the inner wall of the chamber；J：The broken fiber layer scattered pollen grains；ppc：Primary peripheral cells；psc：Primary sporogenous cell；ssc：Secondary sporogenic cell；epi：Epidermis；end：Endothecium；ml：Middle layer；tap：Tapetum

图2 壶瓶枣小孢子和雄配子体的发育过程 A：箭头所指为染色体移向两级；B：箭头所指为二分体；C：箭头所指为二分体至四分体分裂；D：箭头所指为四分体；E：小孢子从四分体释放；F：小孢子单核居中期；G：小孢子单核靠边期；H：生殖细胞形成期；I：成熟花粉粒；J：成熟花粉粒具3个萌发孔。vc：营养细胞；gc：生殖细胞；pa：萌发孔

Fig. 2 The development process of microspore and male gametophyte in Ziziphus jujuba‘Hupingzao’ A：The arrow refers to the chromosome moving to two levels；B：The arrow refers to the dyad；C：The arrow refers to the dyad to tetrad splitting；D：The arrow refers to a tetrad；E：Microspore released from tetrads；F：Microspore monokaryon in middle stage；G：Microspore mononuclear marginal stage；H：The period of germ cell formation；I：Matured pollen grain；J：Mature pollen grains with 3 pollen aperture. vc：Vegetative cell；gc：Germ cell；pa：Pollen aperture

图3 壶瓶枣大孢子的发生过程 A ~ E：倒生胚珠产生过程；F、G：箭头所指为二分体产生过程；H：箭头所指为四分体时期；I：箭头所指为四分体退化；J：箭头所指为大孢子。fu：珠柄；in：珠被；asc：孢原细胞；mea：大孢子

Fig. 3 Macrosporogenesis process of Ziziphus jujuba‘Hupingzao’ A-E：The process of inverted ovule production；F，G：The arrow refers to the process of dyad generation；H：The arrow refers to the tetrad period；I：The arrow refers to the tetrad degeneration；J：The arrow refers to the megaspore. fu：Funicle；in：Integument；asc：Archesporial cell；mea：Megaspore

图4 壶瓶枣雌配子体的发育过程 A：箭头所指为二核胚囊；B：箭头所指为二核胚囊靠近合点端的一个细胞；C：箭头所指为二核胚囊靠近珠孔端的一个细胞；D：箭头所指为合点端的核分裂成两个细胞；E：箭头所指为珠孔端的核分裂成两个细胞；F ~ J：7细胞8核胚囊成熟切片（F：箭头所指为中央细胞；G-H：箭头所指为卵器；I：箭头所指为中央细胞和卵器；J：箭头所指为反足细胞）。cc：中央细胞；ec：卵细胞；sy：助细胞；ea：卵器；ant：反足细胞

Fig. 4 Development process of female gametophyte in Ziziphus jujuba‘Hupingzao’ A：The arrow refers to the two-nuclear embryo sac；B：The arrow refers to a cell near the chalazal end of the binuclear embryo sac；C：The arrow refers to a cell near the micropylar end of the binuclear embryo sac；D：The arrow refers to the nuclear division of the chalazal end into two cells；E：The arrow indicates that the nucleus at the end of the micropylar divides into two cells；F-J：7-cell 8-nuclear embryo sac mature section（F：The arrow refers to the central cell；G，H：The arrow refers to the egg apparatus；I：The arrow refers to the central cell and egg apparatus；J：The arrow refers to antipodal cells）. cc：Central cell；ec：Egg cell； sy：Synergist；ea：Egg apparatus；ant：Antipodal cell

图5 壶瓶枣花的发育过程 se：萼片；st：柱头；an：花药；fi：花丝

Fig. 5 Development process of flower in Ziziphus jujuba‘Hupingzao’ se：Sepal；st：Stigma；an：Anther；fi：Filament

表1 壶瓶枣不同发育时期对应的花形态特征和雌雄蕊发育阶段

Table 1 Floral morphology and developmental stages of pistils and stamens at different growth periods in Ziziphus jujuba‘Hupingzao’

采样时期/（M-D） Sampling stages	花形态特征 Floral morphology	雄蕊发育阶段 Stamen developmental stage	雌蕊发育阶段 Pistil development stage
05-15—05-16	嫩绿色，纵横比0.87 Light green，aspect ratio 0.87	花药原始体 Anther primordium	未分化 Undifferentiation
05-17—05-18	嫩绿色，纵横比0.86 Light green，aspect ratio 0.86	孢原细胞分裂 Archesporial cell division	未分化 Undifferentiation
05-19—05-20	颜色逐渐加深，纵横比0.88 Color gradually deepened，aspect ratio 0.88	孢原细胞平周分裂 Periclinal division of archesporial cells	未分化 Undifferentiation
05-21—05-22	深绿色，纵横比0.84 Dark green，aspect ratio 0.84	花药壁分化，次生造孢细胞形成 Anther wall differentiation，formation of secondary sporogenous cell	未分化 Undifferentiation
05-23—05-24	深绿色，蕾扁期，纵横比0.82 Dark green，bud stage，aspect ratio 0.82	小孢子母细胞形成 Formation of microspore mother cell	未分化 Undifferentiation
05-25—05-26	深绿色，蕾膨大，纵横比0.79 Dark green，bud enlargement，aspect ratio 0.79	小孢子母细胞减数分裂 Microspore mother cell meiosis	胚珠原基 Ovule primordium
05-27—05-28	深绿色，蕾膨大，纵横比0.75 Dark green，bud enlargement，aspect ratio 0.75	小孢子四分体时期 Microspore tetrad stage	孢原细胞造孢细胞发育 Development of archesporial cells into sporogenous cell
05-29—05-30	深绿色，蕾胖期，纵横比0.79 Dark green，bud fat stage，aspect ratio 0.79	小孢子单核居中期 Microspore monokaryon at central position stage	大孢子母细胞形成 Formation of megaspore mother cells
05-31—06-01	黄绿色，纵横比0.69 Yellow green，aspect ratio 0.69	小孢子单核靠边期 Microspore monokaryon at peripheral position stage	二分体—四分体时期 Dyad-tetrad stage
06-02—06-03	黄绿色，纵横比0.74 Yellow green，aspect ratio 0.74	二细胞花粉发育期 Two-celled pollen grain development stage	四分体—单核胚囊 Tetrad- uninucleate embryo sac
06-04—06-05	黄绿色，纵横比0.70 Yellow green，aspect ratio 0.70	二细胞花粉成熟期 Two-celled pollen grain maturation stage	单核胚囊—二核胚囊 Uninucleate embryo sac- binucleate embryo sac
06-06—06-07	黄色，蕾黄期，部分花开，纵横比0.71 Yellow，bud yellow stage，part of the flower blooming，aspect ratio 0.71	成熟花粉粒形成 Mature pollen grain formation	二核胚囊—四核胚囊 Binucleate embryo sac-four nucleate embryo sac
06-08—06-09	黄色，花开放，纵横比0.76 Yellow，flower open，aspect ratio 0.76	成熟花粉粒，花药部分开裂 Mature pollen grain，partial anther dehiscence	四核胚囊—八核胚囊 Four nucleate embryo sac- eight nucleate embryo sac

表1 壶瓶枣不同发育时期对应的花形态特征和雌雄蕊发育阶段

Table 1 Floral morphology and developmental stages of pistils and stamens at different growth periods in Ziziphus jujuba‘Hupingzao’

采样时期/（M-D） Sampling stages	花形态特征 Floral morphology	雄蕊发育阶段 Stamen developmental stage	雌蕊发育阶段 Pistil development stage
05-15—05-16	嫩绿色，纵横比0.87 Light green，aspect ratio 0.87	花药原始体 Anther primordium	未分化 Undifferentiation
05-17—05-18	嫩绿色，纵横比0.86 Light green，aspect ratio 0.86	孢原细胞分裂 Archesporial cell division	未分化 Undifferentiation
05-19—05-20	颜色逐渐加深，纵横比0.88 Color gradually deepened，aspect ratio 0.88	孢原细胞平周分裂 Periclinal division of archesporial cells	未分化 Undifferentiation
05-21—05-22	深绿色，纵横比0.84 Dark green，aspect ratio 0.84	花药壁分化，次生造孢细胞形成 Anther wall differentiation，formation of secondary sporogenous cell	未分化 Undifferentiation
05-23—05-24	深绿色，蕾扁期，纵横比0.82 Dark green，bud stage，aspect ratio 0.82	小孢子母细胞形成 Formation of microspore mother cell	未分化 Undifferentiation
05-25—05-26	深绿色，蕾膨大，纵横比0.79 Dark green，bud enlargement，aspect ratio 0.79	小孢子母细胞减数分裂 Microspore mother cell meiosis	胚珠原基 Ovule primordium
05-27—05-28	深绿色，蕾膨大，纵横比0.75 Dark green，bud enlargement，aspect ratio 0.75	小孢子四分体时期 Microspore tetrad stage	孢原细胞造孢细胞发育 Development of archesporial cells into sporogenous cell
05-29—05-30	深绿色，蕾胖期，纵横比0.79 Dark green，bud fat stage，aspect ratio 0.79	小孢子单核居中期 Microspore monokaryon at central position stage	大孢子母细胞形成 Formation of megaspore mother cells
05-31—06-01	黄绿色，纵横比0.69 Yellow green，aspect ratio 0.69	小孢子单核靠边期 Microspore monokaryon at peripheral position stage	二分体—四分体时期 Dyad-tetrad stage
06-02—06-03	黄绿色，纵横比0.74 Yellow green，aspect ratio 0.74	二细胞花粉发育期 Two-celled pollen grain development stage	四分体—单核胚囊 Tetrad- uninucleate embryo sac
06-04—06-05	黄绿色，纵横比0.70 Yellow green，aspect ratio 0.70	二细胞花粉成熟期 Two-celled pollen grain maturation stage	单核胚囊—二核胚囊 Uninucleate embryo sac- binucleate embryo sac
06-06—06-07	黄色，蕾黄期，部分花开，纵横比0.71 Yellow，bud yellow stage，part of the flower blooming，aspect ratio 0.71	成熟花粉粒形成 Mature pollen grain formation	二核胚囊—四核胚囊 Binucleate embryo sac-four nucleate embryo sac
06-08—06-09	黄色，花开放，纵横比0.76 Yellow，flower open，aspect ratio 0.76	成熟花粉粒，花药部分开裂 Mature pollen grain，partial anther dehiscence	四核胚囊—八核胚囊 Four nucleate embryo sac- eight nucleate embryo sac

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