木麻黄生殖生物学研究
摘要
木麻黄(Casuarinaceae)杂交育种是木麻黄遗传改良的重要手段。杂种优势的利用,在现代育种中成果突出,有性制种无性繁殖能大幅度提高木麻黄人工林产量及对不良环境的适应能力和抗性。本试验主要研究了木麻黄的生殖生物学习性,对木麻黄的雌雄花的结构、花粉活力、花粉贮藏时间及贮藏条件、花粉的形态特征、木麻黄传粉受精后胚胎发育的不同阶段和木麻黄种子抗盐性,进行了较深入的调查、探索和分析研究,得出了如下结论：
1木麻黄的雌雄花的结构
木麻黄花单性,雌雄同株或异株；雌花是柔荑花序。雄花序穗状,雄花序长约1-5cm,几无总花梗；有1枚雄蕊和4个苞片,花药为二心房四心室；雌花序为球形或头状,顶生于短的侧枝上,较雄花序短而宽；雌花1枚苞片和2枚小苞片腋生,无花被；雌蕊由2枚心皮组成。子房上位,初为2室,因后位退化而成为单室,花柱短,有2条通常为红色的线形柱头。
2花粉活力测定
从试验结果可以看出,不同蔗糖、硼酸浓度对木麻黄花粉萌发有显著的影响,通过试验筛选出花粉萌发的最佳蔗糖和硼酸浓度。15%的蔗糖浓度是木麻黄花粉萌发的最佳浓度；在15%蔗糖培养基上添加硼酸能够显著促进木麻黄花粉萌发,250 mg kg-1硼酸浓度是木麻黄花粉萌发的最佳浓度；研究也证明添加了琼脂的固体培养基上更有利于木麻黄花粉萌发。木麻黄花粉的发芽率偏低,活力较低,即使在最优组合的培养条件下最高的发芽率也只有13.16%。
3木麻黄花粉的贮藏条件及时间
研究表明,木麻黄花粉萌发率在各种贮藏温度下都随贮藏时间的延长而下降,但在不同的贮藏温度下花粉萌发率的下降速度差异很大。在室温条件下,花粉在贮藏3天的萌发率只有0.79%,贮藏7天的花粉已经没有活力；在冰箱4℃冷藏的条件下,木麻黄花粉贮藏3天的萌发率11.20%,贮藏7天的萌发率有8.47%,贮藏15天的萌发率降低为4.79%,贮藏30天后的萌发率只有2.33%；而在冰箱-20℃冷冻的条件下,木麻黄花粉贮藏3天的萌发率达13.16%,7天的萌发率为10.32%,贮藏15天后下降为9.86%,贮藏30天后萌发率仍有8.32%。这说明木麻黄花粉在室温的条件下贮藏时间很短,3天后绝大部分花粉失去了活力。试验也证明低温条件能非常有效地保持木麻黄花粉的生活力,如贮藏30天的花粉萌发率比贮藏7天的萌发率仅降低了2.0%。通过花粉的贮藏试验,探讨木麻黄花粉在不同贮藏条件下生活力的变化,为花粉的长时间贮藏和远距离运输提供科学依据。
4木麻黄花粉形态特征的观测
木麻黄花粉在赤道面观呈圆球形或近球形,有三个萌发孔,均等的分布在花粉球表面。从各图版高倍图像可知,所观察的3种木麻黄,短枝木麻黄(C. equisetifolia),细枝木麻黄(C. cunninghamiana Miq.)和粗枝木麻黄(C. glauca Sieber ex Sprengel)的花粉粒的表面纹饰的基本类型都具有刺突,但不同种间刺突的分布密度不均,表面的粗糙程度和有无褶皱以及花粉大小也有差异。短枝木麻的花粉近球形,直径在28.8-39.2μm之间,表面粗糙,有褶皱,刺突分布密集。细枝木麻黄的花粉是圆球形,直径在28.6-38.6μm之间,表面粗糙,有褶皱,但是刺突的分布密度不同无性系间还有差异；粗枝木麻黄的花粉圆球形,直径在29.8-46.1μm之间,表面光滑程度不同无性系间有差异,有的无性系有褶皱,有的无褶皱,刺突分布疏松。
5木麻黄胚胎发育的不同阶段
试验对木麻黄的胚发育做了简单了解,对不同时期的胚形态进行了初步的学习和研究,但是由于采样时间的问题,没有系统的掌握木麻黄胚发育的整个完整时期,只能简单地了解了木麻黄胚珠的结构,整个胚的发育史还有待进一步的研究学习。
6木麻黄种子抗盐性
实验研究了在不同浓度的NaCl胁迫下,山地木麻黄(C.montana Miq)、细枝木麻黄和粗枝木麻黄种子萌发和幼苗的生长状况。结果表明,随着NaCl浓度的增加,3个树种的种子萌发率、活力指数、发芽势以及幼苗的苗高和根长均呈下降趋势,结果表明,木麻黄种子在盐胁迫下,发芽率、发芽指数、活力指数都随着盐浓度的升高而降低,萌发后获得的小幼苗的早期生长受到一定的抑制；随着NaCl浓度的增大,对种子萌发、幼苗生长的抑制作用越强,不同NaCl浓度的影响差异显著；当NaCl浓度为0.3%时,对3组木麻黄种子的发芽率、发芽指数、活力指数以及幼苗高、根长、鲜重等与对照相比下降幅度较大,受到明显抑制,而不同种之间的木麻黄种子的发芽率差异显著。
The crossbreeding of Casuarinaceae is a very important method of Casuarinaceae genetic improvement. The Use of hybrids advantage is particularly prominent in modern breeding, sexual hybrid seed production can greatly improve the production of plantations,the adaptability and resistance of bad environment. This experiment mainly studied the reproductive and biological of Casuarinaceae, the structure of male and female flowers of Casuarinaceae、pollen viability、the storage time and condition of pollen、he morphology of pollen、different stages of embryo development after pollination and fertilization, and conclusions as follow: 1.The structure of male and female flowers of Casuarinaceae
The flower of Casuarinaceae is unisexual, it is monoecious or dioecious; catkin. Male inflorescenceis like panicle, it is 1-5cm long, almost no peduncle; 1 stamen and 4 bracts, anther with 2 atrials and 4 ventriculars; female inflorescence is like a ball, and it is on the short lateral branch,shorter wider than male inflorescenceis; female flower has 1 bract and 2 little axillary bracts, no perianth; pistil is formed by 2 carpels. superior ovary,2 rooms initially, then back part degraded into single room, the style is short, and 2 red lineshape stigma. 2. The determination of pollen viability
The test results shows that,ifferent concentration of sucrose and boric acid has obvious influence to the germination of pollen. screening the best concentration of sucrose and boric acid for the pollen germination.15% of concentration of sucrose is the best concentration for the germination of Casuarinaceae pollen; adding boric acid to the medium with 15% of concentration of sucrose can improve the pollen germination,250 mgkg-1 of boric acid is the best concentration for the germination Casuarinaceae pollen; the research also proved that the solid medium with agar is better for the germination of Casuarinaceae pollen.The germination of Casuarinaceae pollen is low, the activity is low too, the highest germination is only 13.16% even under the culture condition with the best combination. 3.The storage time and condition of pollen
The research shows that, the germination rate of Casuarinaceae pollen is decreased with the prolongation of the storage time under many kinds of storage time but the difference of descending speed is great under different storage temperature under the condition of room temperature, the germination rate of pollen is only 0.79% after 3days storage, and the pollen has no activity after 7 days storage; under the condition of 4℃in fridge, the germination rate of pollen is only 11.20% after 3days storage,only 8.47%, and it decreased to4.79% after 15 days storage,2.33% after 30 days; but under the condition of-20℃in fridge, the germination rate of pollen is only 13.16% after 3days storage,10.32% afer 7 days storage,9.86% after 15 days storage, and 8.32% after 30 days storage still. It showed that the storage time of Casuarinaceae pollen is very short under the condition of room temperature, most pollen lost its activity after 3 days.The experiment also proved that low temperature can keep the activity of Casuarinaceae pollen effectively, the germination rate of pollen after 30 days storage is 2.0% lower than the one after 7 days.In the experiment of pollen storage,studying the change under different storage condition can provide scientific basis for long time storage and transform of the pollen.
4. The observation of the morphological characters of Casuarinaceae pollen
The equatorial view of Casuarinaceae is coccoid form or near spherical,and it has 3 apertures, they distribut on the face of the pollen.We can see from all the pictures, the tree kinds of Casuarinaceae, C. equisetifolia, C. cunninghamiana Miq.and C. glauca Sieber ex Sprengel, the face ornamentation of the pollen all have spikes,but the density of different kinds of spikes is uneven, the rough extent and folds and the size of the pollen is different either.C. equisetifolia: The pollen of C.equisetifolia is near spherical, the ameter is between 28.8-39.2μm, it has roughface and folds, the distributionof of spikes is densely. The pollen of C. cunninghamiana Miq is coccoid form, the ameter is between28.6-38.6μm, it has rough face and folds, the distributionof of spikes is different and it has difference between different clones; The pollen of C. glauca Sieber ex Sprengel is also coccoid form, the ameter is between 27.7-35.6μmm, it has rough face and folds, the smooth degree is different and it has difference between different clones, some has folds while some has no folds, the distributionof of spikes is densely.
5.Different stages of embryo development of Casuarinaceae
The experiment did a simple understand of embryo development, and initial study and research of morphological embryo in different period,but because the sampling time problem, we did not master the hole period of Casuarinaceae embryo development systematicly, and just study the structure of Casuarinaceae ovule simply, so we need to study the hole life cycle of the embryo further more.
6. Seeds of Casuarina ability to resist salt
The experiment has studied the seed germination and the growth conditionof seedling of C.montana Miq、C. cunninghamiana Miq.and C. glauca Sieber ex Sprengel under the condition of different concentration of NaCl. And the result showed that the germination rate、vigor index、germination energy、the seedling heightand the root length of the 3 kinds of trees is in a decreased trend. Under the salt stress, the germination rate、germination index、vigor index of the Casuarinaceae seed is decreased with the increase of the concentration of salt, the early growth of seedlings after germination is inhibited; the inhibition of seed germination and seedling growth is great with the increase of the concentration of NaCl; when the concentration of NaCl is 0.3%, the decrease amplitude of germination rate、germination index、vigor index、germination energy、the seedling height、the root length and fresh weight were larger than the normal,they were inhibited greatly, but the germination rate between different kinds of Casuarinaceae seed is significant different.
1木麻黄的雌雄花的结构
木麻黄花单性,雌雄同株或异株；雌花是柔荑花序。雄花序穗状,雄花序长约1-5cm,几无总花梗；有1枚雄蕊和4个苞片,花药为二心房四心室；雌花序为球形或头状,顶生于短的侧枝上,较雄花序短而宽；雌花1枚苞片和2枚小苞片腋生,无花被；雌蕊由2枚心皮组成。子房上位,初为2室,因后位退化而成为单室,花柱短,有2条通常为红色的线形柱头。
2花粉活力测定
从试验结果可以看出,不同蔗糖、硼酸浓度对木麻黄花粉萌发有显著的影响,通过试验筛选出花粉萌发的最佳蔗糖和硼酸浓度。15%的蔗糖浓度是木麻黄花粉萌发的最佳浓度；在15%蔗糖培养基上添加硼酸能够显著促进木麻黄花粉萌发,250 mg kg-1硼酸浓度是木麻黄花粉萌发的最佳浓度；研究也证明添加了琼脂的固体培养基上更有利于木麻黄花粉萌发。木麻黄花粉的发芽率偏低,活力较低,即使在最优组合的培养条件下最高的发芽率也只有13.16%。
3木麻黄花粉的贮藏条件及时间
研究表明,木麻黄花粉萌发率在各种贮藏温度下都随贮藏时间的延长而下降,但在不同的贮藏温度下花粉萌发率的下降速度差异很大。在室温条件下,花粉在贮藏3天的萌发率只有0.79%,贮藏7天的花粉已经没有活力；在冰箱4℃冷藏的条件下,木麻黄花粉贮藏3天的萌发率11.20%,贮藏7天的萌发率有8.47%,贮藏15天的萌发率降低为4.79%,贮藏30天后的萌发率只有2.33%；而在冰箱-20℃冷冻的条件下,木麻黄花粉贮藏3天的萌发率达13.16%,7天的萌发率为10.32%,贮藏15天后下降为9.86%,贮藏30天后萌发率仍有8.32%。这说明木麻黄花粉在室温的条件下贮藏时间很短,3天后绝大部分花粉失去了活力。试验也证明低温条件能非常有效地保持木麻黄花粉的生活力,如贮藏30天的花粉萌发率比贮藏7天的萌发率仅降低了2.0%。通过花粉的贮藏试验,探讨木麻黄花粉在不同贮藏条件下生活力的变化,为花粉的长时间贮藏和远距离运输提供科学依据。
4木麻黄花粉形态特征的观测
木麻黄花粉在赤道面观呈圆球形或近球形,有三个萌发孔,均等的分布在花粉球表面。从各图版高倍图像可知,所观察的3种木麻黄,短枝木麻黄(C. equisetifolia),细枝木麻黄(C. cunninghamiana Miq.)和粗枝木麻黄(C. glauca Sieber ex Sprengel)的花粉粒的表面纹饰的基本类型都具有刺突,但不同种间刺突的分布密度不均,表面的粗糙程度和有无褶皱以及花粉大小也有差异。短枝木麻的花粉近球形,直径在28.8-39.2μm之间,表面粗糙,有褶皱,刺突分布密集。细枝木麻黄的花粉是圆球形,直径在28.6-38.6μm之间,表面粗糙,有褶皱,但是刺突的分布密度不同无性系间还有差异；粗枝木麻黄的花粉圆球形,直径在29.8-46.1μm之间,表面光滑程度不同无性系间有差异,有的无性系有褶皱,有的无褶皱,刺突分布疏松。
5木麻黄胚胎发育的不同阶段
试验对木麻黄的胚发育做了简单了解,对不同时期的胚形态进行了初步的学习和研究,但是由于采样时间的问题,没有系统的掌握木麻黄胚发育的整个完整时期,只能简单地了解了木麻黄胚珠的结构,整个胚的发育史还有待进一步的研究学习。
6木麻黄种子抗盐性
实验研究了在不同浓度的NaCl胁迫下,山地木麻黄(C.montana Miq)、细枝木麻黄和粗枝木麻黄种子萌发和幼苗的生长状况。结果表明,随着NaCl浓度的增加,3个树种的种子萌发率、活力指数、发芽势以及幼苗的苗高和根长均呈下降趋势,结果表明,木麻黄种子在盐胁迫下,发芽率、发芽指数、活力指数都随着盐浓度的升高而降低,萌发后获得的小幼苗的早期生长受到一定的抑制；随着NaCl浓度的增大,对种子萌发、幼苗生长的抑制作用越强,不同NaCl浓度的影响差异显著；当NaCl浓度为0.3%时,对3组木麻黄种子的发芽率、发芽指数、活力指数以及幼苗高、根长、鲜重等与对照相比下降幅度较大,受到明显抑制,而不同种之间的木麻黄种子的发芽率差异显著。
The crossbreeding of Casuarinaceae is a very important method of Casuarinaceae genetic improvement. The Use of hybrids advantage is particularly prominent in modern breeding, sexual hybrid seed production can greatly improve the production of plantations,the adaptability and resistance of bad environment. This experiment mainly studied the reproductive and biological of Casuarinaceae, the structure of male and female flowers of Casuarinaceae、pollen viability、the storage time and condition of pollen、he morphology of pollen、different stages of embryo development after pollination and fertilization, and conclusions as follow: 1.The structure of male and female flowers of Casuarinaceae
The flower of Casuarinaceae is unisexual, it is monoecious or dioecious; catkin. Male inflorescenceis like panicle, it is 1-5cm long, almost no peduncle; 1 stamen and 4 bracts, anther with 2 atrials and 4 ventriculars; female inflorescence is like a ball, and it is on the short lateral branch,shorter wider than male inflorescenceis; female flower has 1 bract and 2 little axillary bracts, no perianth; pistil is formed by 2 carpels. superior ovary,2 rooms initially, then back part degraded into single room, the style is short, and 2 red lineshape stigma. 2. The determination of pollen viability
The test results shows that,ifferent concentration of sucrose and boric acid has obvious influence to the germination of pollen. screening the best concentration of sucrose and boric acid for the pollen germination.15% of concentration of sucrose is the best concentration for the germination of Casuarinaceae pollen; adding boric acid to the medium with 15% of concentration of sucrose can improve the pollen germination,250 mgkg-1 of boric acid is the best concentration for the germination Casuarinaceae pollen; the research also proved that the solid medium with agar is better for the germination of Casuarinaceae pollen.The germination of Casuarinaceae pollen is low, the activity is low too, the highest germination is only 13.16% even under the culture condition with the best combination. 3.The storage time and condition of pollen
The research shows that, the germination rate of Casuarinaceae pollen is decreased with the prolongation of the storage time under many kinds of storage time but the difference of descending speed is great under different storage temperature under the condition of room temperature, the germination rate of pollen is only 0.79% after 3days storage, and the pollen has no activity after 7 days storage; under the condition of 4℃in fridge, the germination rate of pollen is only 11.20% after 3days storage,only 8.47%, and it decreased to4.79% after 15 days storage,2.33% after 30 days; but under the condition of-20℃in fridge, the germination rate of pollen is only 13.16% after 3days storage,10.32% afer 7 days storage,9.86% after 15 days storage, and 8.32% after 30 days storage still. It showed that the storage time of Casuarinaceae pollen is very short under the condition of room temperature, most pollen lost its activity after 3 days.The experiment also proved that low temperature can keep the activity of Casuarinaceae pollen effectively, the germination rate of pollen after 30 days storage is 2.0% lower than the one after 7 days.In the experiment of pollen storage,studying the change under different storage condition can provide scientific basis for long time storage and transform of the pollen.
4. The observation of the morphological characters of Casuarinaceae pollen
The equatorial view of Casuarinaceae is coccoid form or near spherical,and it has 3 apertures, they distribut on the face of the pollen.We can see from all the pictures, the tree kinds of Casuarinaceae, C. equisetifolia, C. cunninghamiana Miq.and C. glauca Sieber ex Sprengel, the face ornamentation of the pollen all have spikes,but the density of different kinds of spikes is uneven, the rough extent and folds and the size of the pollen is different either.C. equisetifolia: The pollen of C.equisetifolia is near spherical, the ameter is between 28.8-39.2μm, it has roughface and folds, the distributionof of spikes is densely. The pollen of C. cunninghamiana Miq is coccoid form, the ameter is between28.6-38.6μm, it has rough face and folds, the distributionof of spikes is different and it has difference between different clones; The pollen of C. glauca Sieber ex Sprengel is also coccoid form, the ameter is between 27.7-35.6μmm, it has rough face and folds, the smooth degree is different and it has difference between different clones, some has folds while some has no folds, the distributionof of spikes is densely.
5.Different stages of embryo development of Casuarinaceae
The experiment did a simple understand of embryo development, and initial study and research of morphological embryo in different period,but because the sampling time problem, we did not master the hole period of Casuarinaceae embryo development systematicly, and just study the structure of Casuarinaceae ovule simply, so we need to study the hole life cycle of the embryo further more.
6. Seeds of Casuarina ability to resist salt
The experiment has studied the seed germination and the growth conditionof seedling of C.montana Miq、C. cunninghamiana Miq.and C. glauca Sieber ex Sprengel under the condition of different concentration of NaCl. And the result showed that the germination rate、vigor index、germination energy、the seedling heightand the root length of the 3 kinds of trees is in a decreased trend. Under the salt stress, the germination rate、germination index、vigor index of the Casuarinaceae seed is decreased with the increase of the concentration of salt, the early growth of seedlings after germination is inhibited; the inhibition of seed germination and seedling growth is great with the increase of the concentration of NaCl; when the concentration of NaCl is 0.3%, the decrease amplitude of germination rate、germination index、vigor index、germination energy、the seedling height、the root length and fresh weight were larger than the normal,they were inhibited greatly, but the germination rate between different kinds of Casuarinaceae seed is significant different.
引文
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