平邑甜茶无融合生殖发育与遗传特性分析及相关基因的功能鉴定
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摘要
无融合生殖是发育生物学领域的重要科学问题。与有性种子后代相比，无融合生殖后代的遗传背景与母本一致，不发生性状分离。在水稻和玉米等作物育种中，无融合生殖可以固定杂种优势；在苹果、柑橘、马铃薯和甘薯等无性繁殖植物中，无融合生殖产生无性种子，克服无性繁殖的不便，并能避免长期营养繁殖造成的亲本退化。鉴于无融合生殖在农业生产中的巨大应用潜力，并受到植物学家和育种学家的广泛关注，对无融合生殖进行相关研究不仅具有重要的理论价值，而且具有潜在的实践意义。
本文以兼性无融合生殖的野生苹果资源平邑甜茶（Malus hupehensis Redh. var.pingyiensis）及其有性杂交群体为试材，探讨苹果属无融合生殖植物的发育和遗传特性及其分子机理。主要研究结果如下：
1.平邑甜茶具有高度单性生殖和2n雌配子形成能力
平邑甜茶胚囊由胚珠体细胞发育而来，并存在多胚囊现象，属于无孢子无融合生殖类型。通过连续3年的调查统计，发现平邑甜茶的单性生殖能力平均为94%，2n雌配子形成能力平均为95%，说明平邑甜茶具有高度的单性生殖能力和无融合非减数分裂能力。
2.平邑甜茶等苹果属无融合生殖资源的2n胚囊通过发育迟滞策略逃避受精
平邑甜茶的2n胚囊能够正常受精，但由于发育滞后，当其发育成熟时，柱头已经衰老并失去授粉能力，从而逃避了受精过程，最终通过单性生殖形成无融合生殖种子，我们把这种现象称之为“FLD”。FLD现象在小金海棠和沙金海棠等其它无融合生殖苹果资源中也普遍存在，表明胚囊发育迟滞是苹果属无融合生殖过程的关键发育事件。
3.缩短柱头寿命提高无融合生殖率
苹果属无融合生殖材料大部分进行的是兼性无融合生殖，自然授粉下可形成有性种子，而通过喷施石硫合剂可以缩短柱头寿命，阻止授粉受精，从而降低杂种率，提高无融合生殖率。
4.无融合非减数分裂和单性生殖独立遗传并共同决定无融合生殖
利用平邑甜茶与舞美的杂交后代对无融合生殖特性进行遗传分析，结果表明，2n雌配子形成能力与单性生殖能力不偶联，是相互独立的发育事件，并且两者的乘积与无融合生殖率和FLD现象正相关。所以，平邑甜茶的无融合生殖由单性生殖和2n雌配子形成两个发育事件组成，而胚囊发育迟滞现象是反映无融合生殖能力的关键早期发育事件。
5.无融合生殖相关基因的筛选和克隆
平邑甜茶与舞美的杂交后代群体，根据无融合生殖能力的高低分为高-低无融合非减数分裂群体和高-低单性生殖群体，这两组群体的表达谱分析，筛选表达差异的无融合生殖相关基因。根据筛选结果选择MhFIE和MhMSI1两个基因，克隆测序后发现二者均含有WD40保守区域，GFP融合蛋白分析发现，它们均定位于细胞核。
6. MhFIE和MhMSI1基因的功能鉴定
为了鉴定MhFIE和MhMSI1基因的生物学功能，将35S启动子驱动的MhFIE和MhMSI1基因分别遗传转化拟南芥、番茄和苹果，实现两个基因在拟南芥和番茄等植物中的异位表达，与对照相比，转基因株系的营养生长和生殖器官发育均表现出异常，表明MhFIE和MhMSI1基因在植物营养生长与生殖生长中均具有重要功能。
Apomixis is an attractive trait in developmental biology. Compared with the sexualoffspring, progeny of apomixis are large genetically uniform and without character separation.In the rice and corn breeding, apomixis can fix heterosis. In addition, apple, orange, potatoesand sweet potatoes are typically propagated vegetatively. if apomixis is introduced to them,they would produced asexual seeds which can avoid inconvenience in propagation anddegradation in long-term vegetative propagation. Therefore, apomixis has great potentialapplication in agricultural production. Because of its huge potential utilization inhybridization breeding, both botanists and breeders are interested in elucidating the molecularmechanism underlying apomixis.
In this study, developmental and genetic characters were observed and investigated inapomictic tea crabapple. Apomixis-related genes were molecular cloned and functionallycharacterized. The main results are shown as follows:
1. Tea crabapple shows high capacity of parthenogenesis and apomeiosis
Stomatic cells developed into one or more embryo sacs in the ovules of tea crabapple.Aposporous embryo sac initiated development at tetrad stage. Investigations throughcontinuous three years showed that the percentages of parthenogenesis and2n gamete were94%and95%, respectively, in tea crabapple, indicating that it exhibits high capacity toperform parthenogenesis and apomeiosis.
2.2n embryo sacs escape fertilization due to developmental delay in apomictic apple species
Embryo sacs of tea crabapple showed developmental delay,which was also found both inother two apomictic wild apple species M. Sargentii and M. Xiaojinensis. When embryo sacbecame mature, the stigma turned too senescent to accept pollens. Finally, asexual seedsformed directly from2n embryo sacs without fertilization. This phenomenon was called FLD,which is an early developmental event crucial for apomixis.
3. Apomictic reproduction is enhanced by shortening pistil longevity
Most of apomictic apple species perform facultative apomixes, i.e. they produce sexualseeds to certain great in addition to apomictic ones. Apomictic seed formation was enhancedby spraying lime sulphur in order to shorten pistil longevity and avoid pollination.
4. Apomeiosis and parthenogenesis are genetically independent but collaborate to determineapomixis
Genetical analysis showed that apomeiosis and parthenogenesis are geneticallyindependent to each other. They were uncoupled in the hybrid population obtained from a cross between tea crabapple and Maypole. But their multiplication was positively related tothe apomictic capacity and FLD phenomenon. Therefore, apomeiosis and parthenogenesisdetermine apomictic capacity, and FLD reflect the ability of apomixis in apomictic applespecies.
5. Screening and cloning of apomixis-related genes
According to the different ability of apomeiosis and parthenogenesis, hybrid offspringpopulation was divided into four subpopulations. Apomixis genes were screened through anexpression analysis using cDNA libraries of4subpopulations. Depending on the screening,two genes MhFIE and MhMSI1were cloned. Both predicted protein contained WD domainswhich are highly conserved in various plant species. Furthermore, GFP fusion analysisdemonstrated that both MhFIE and MhMSI1proteins were subcellurly localized in nucleus.
6. Functional characterization of MhFIE and MhMSI1genes
To further characterize the functions of MhFIE and MhMSI1in planta, the constructscontaining MhFIE and MhMSI1driven by35S promoter were genetically transformed intoarabidopsis, tomato and apple, respectively. In transgenic Arabidopsis and tomato lines,ectopic expression of MhFIE and MhMSI1brought about developmental alterations both invegetative and reproductive growth, indicating that the functions of both genes are importantfor normal growth in higher plants.
本文以兼性无融合生殖的野生苹果资源平邑甜茶（Malus hupehensis Redh. var.pingyiensis）及其有性杂交群体为试材，探讨苹果属无融合生殖植物的发育和遗传特性及其分子机理。主要研究结果如下：
1.平邑甜茶具有高度单性生殖和2n雌配子形成能力
平邑甜茶胚囊由胚珠体细胞发育而来，并存在多胚囊现象，属于无孢子无融合生殖类型。通过连续3年的调查统计，发现平邑甜茶的单性生殖能力平均为94%，2n雌配子形成能力平均为95%，说明平邑甜茶具有高度的单性生殖能力和无融合非减数分裂能力。
2.平邑甜茶等苹果属无融合生殖资源的2n胚囊通过发育迟滞策略逃避受精
平邑甜茶的2n胚囊能够正常受精，但由于发育滞后，当其发育成熟时，柱头已经衰老并失去授粉能力，从而逃避了受精过程，最终通过单性生殖形成无融合生殖种子，我们把这种现象称之为“FLD”。FLD现象在小金海棠和沙金海棠等其它无融合生殖苹果资源中也普遍存在，表明胚囊发育迟滞是苹果属无融合生殖过程的关键发育事件。
3.缩短柱头寿命提高无融合生殖率
苹果属无融合生殖材料大部分进行的是兼性无融合生殖，自然授粉下可形成有性种子，而通过喷施石硫合剂可以缩短柱头寿命，阻止授粉受精，从而降低杂种率，提高无融合生殖率。
4.无融合非减数分裂和单性生殖独立遗传并共同决定无融合生殖
利用平邑甜茶与舞美的杂交后代对无融合生殖特性进行遗传分析，结果表明，2n雌配子形成能力与单性生殖能力不偶联，是相互独立的发育事件，并且两者的乘积与无融合生殖率和FLD现象正相关。所以，平邑甜茶的无融合生殖由单性生殖和2n雌配子形成两个发育事件组成，而胚囊发育迟滞现象是反映无融合生殖能力的关键早期发育事件。
5.无融合生殖相关基因的筛选和克隆
平邑甜茶与舞美的杂交后代群体，根据无融合生殖能力的高低分为高-低无融合非减数分裂群体和高-低单性生殖群体，这两组群体的表达谱分析，筛选表达差异的无融合生殖相关基因。根据筛选结果选择MhFIE和MhMSI1两个基因，克隆测序后发现二者均含有WD40保守区域，GFP融合蛋白分析发现，它们均定位于细胞核。
6. MhFIE和MhMSI1基因的功能鉴定
为了鉴定MhFIE和MhMSI1基因的生物学功能，将35S启动子驱动的MhFIE和MhMSI1基因分别遗传转化拟南芥、番茄和苹果，实现两个基因在拟南芥和番茄等植物中的异位表达，与对照相比，转基因株系的营养生长和生殖器官发育均表现出异常，表明MhFIE和MhMSI1基因在植物营养生长与生殖生长中均具有重要功能。
Apomixis is an attractive trait in developmental biology. Compared with the sexualoffspring, progeny of apomixis are large genetically uniform and without character separation.In the rice and corn breeding, apomixis can fix heterosis. In addition, apple, orange, potatoesand sweet potatoes are typically propagated vegetatively. if apomixis is introduced to them,they would produced asexual seeds which can avoid inconvenience in propagation anddegradation in long-term vegetative propagation. Therefore, apomixis has great potentialapplication in agricultural production. Because of its huge potential utilization inhybridization breeding, both botanists and breeders are interested in elucidating the molecularmechanism underlying apomixis.
In this study, developmental and genetic characters were observed and investigated inapomictic tea crabapple. Apomixis-related genes were molecular cloned and functionallycharacterized. The main results are shown as follows:
1. Tea crabapple shows high capacity of parthenogenesis and apomeiosis
Stomatic cells developed into one or more embryo sacs in the ovules of tea crabapple.Aposporous embryo sac initiated development at tetrad stage. Investigations throughcontinuous three years showed that the percentages of parthenogenesis and2n gamete were94%and95%, respectively, in tea crabapple, indicating that it exhibits high capacity toperform parthenogenesis and apomeiosis.
2.2n embryo sacs escape fertilization due to developmental delay in apomictic apple species
Embryo sacs of tea crabapple showed developmental delay,which was also found both inother two apomictic wild apple species M. Sargentii and M. Xiaojinensis. When embryo sacbecame mature, the stigma turned too senescent to accept pollens. Finally, asexual seedsformed directly from2n embryo sacs without fertilization. This phenomenon was called FLD,which is an early developmental event crucial for apomixis.
3. Apomictic reproduction is enhanced by shortening pistil longevity
Most of apomictic apple species perform facultative apomixes, i.e. they produce sexualseeds to certain great in addition to apomictic ones. Apomictic seed formation was enhancedby spraying lime sulphur in order to shorten pistil longevity and avoid pollination.
4. Apomeiosis and parthenogenesis are genetically independent but collaborate to determineapomixis
Genetical analysis showed that apomeiosis and parthenogenesis are geneticallyindependent to each other. They were uncoupled in the hybrid population obtained from a cross between tea crabapple and Maypole. But their multiplication was positively related tothe apomictic capacity and FLD phenomenon. Therefore, apomeiosis and parthenogenesisdetermine apomictic capacity, and FLD reflect the ability of apomixis in apomictic applespecies.
5. Screening and cloning of apomixis-related genes
According to the different ability of apomeiosis and parthenogenesis, hybrid offspringpopulation was divided into four subpopulations. Apomixis genes were screened through anexpression analysis using cDNA libraries of4subpopulations. Depending on the screening,two genes MhFIE and MhMSI1were cloned. Both predicted protein contained WD domainswhich are highly conserved in various plant species. Furthermore, GFP fusion analysisdemonstrated that both MhFIE and MhMSI1proteins were subcellurly localized in nucleus.
6. Functional characterization of MhFIE and MhMSI1genes
To further characterize the functions of MhFIE and MhMSI1in planta, the constructscontaining MhFIE and MhMSI1driven by35S promoter were genetically transformed intoarabidopsis, tomato and apple, respectively. In transgenic Arabidopsis and tomato lines,ectopic expression of MhFIE and MhMSI1brought about developmental alterations both invegetative and reproductive growth, indicating that the functions of both genes are importantfor normal growth in higher plants.
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