树莓生殖生物学特性研究
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摘要
树莓为属于蔷薇科(Rosaceae)悬钩子属(Rubus L.)的小浆果果树,按其结果时间可分为夏果型和秋果型红树莓。秋果型红树莓2年生枝条和1年生枝条均可结果。如果只利用其1年生枝条结果冬季可不用埋土,因而可降低生产成本。目前我国仅见关于夏果型树莓的花芽分化的报道,而有关秋果型花芽分化规律的研究还尚未开展,在树莓其它生殖生物学方面的研究也还属空白。因而本研究以夏果型品种奥瑞斯和秋果型品种撒米特为研究试材,采用石蜡切片的方法对其花芽形态分化、大小孢子和雌雄配子体的发育、果实的解剖构造进行了观察,采用TTC法测定常温和低温下贮藏花粉活力的变化,采用扫描电镜的方法观察了树莓的花粉形态,同时对树莓的有效授粉期、柱头可授性、不同授粉方式对座果率的影响,种子不同处理方法对其发芽力的影响进行了研究,主要研究结果如下:
1.树莓花芽形态分化过程可分为花芽未分化期、花芽分化初期、小花分化期三个时期,小花分化期又包括花萼原基、雄蕊原基和大量离生心皮组成的雌蕊原基时期,花瓣原基。在哈尔滨地区,品种撒米特的2年生枝条、1年生枝条和奥瑞斯的2年生枝条花芽分化进程基本一致,分化时间不同。
2.大小孢子发育和雌雄配子体的发育特点为花药壁属于双子叶型,绒毡层为腺质(分泌)绒毡层,小孢子母细胞为多数,减数分裂四分体呈四面体型,小孢子圆形,较小,雄配子体为2-细胞型;蓼型胚囊,每子房内有2个胚珠,其中1个退化,另1个正常发育。倒生胚珠。
3.成熟花粉粒圆型,具有三个萌发沟,花粉外壁呈网状,萌发孔具有孔盖,萌发沟、表层具有小颗粒。
4.采用TTC测定保存在常温下和冰箱保鲜层中的花粉,表明无论在常温下,还是在冰箱保鲜层中保存20天后撒米特的花粉活性仍可达到50%以上,而奥瑞斯的花粉分别降低到38.26%和44.59%。
5.奥瑞斯柱头可授性可持续24小时,撒米特柱头可授性可持续30小时。奥瑞斯有效授粉期为7天,撒米特有效授粉期为4天。奥瑞斯自交结实率、自然结实率和人工授粉结实率分别为:52.73%、84.73%、72.46%;撒米特自交结实率、自然结实率和人工授粉结实率分别为:46.33%、90.62%、79.61%。
6.解剖观察表明树莓果实由果皮和种子两部分组成。果皮自外向内具有三层,外果皮较薄,表面光滑。中果皮由多层薄壁细胞组成,内果皮坚硬,由大量纤维状厚壁细胞组成,果实属于聚合核果类型;种子1枚,由种皮和胚组成,种皮较厚,多层细胞组成,排列紧密。
7.树莓果实的生长曲线表现为双S曲线,在两次速长期之间有1个缓慢生长期,但持续时间短。
Raspberry, small fruit of Rosaceae, Rubus L., has two types according to the time of fruit forming: summer bearing and primocane bearing. Both 2-years and 1-year cane of primocane bearing type can produce fruits and we can use 1-year cane, which need not be buried in winter to reduce the cost. Now, there are some reports about the flower bud differentiation of summer-fruit type Raspberry in China and there is still a blank in the study of Raspberry procreate biology. In this paper, we took summer-fruit type‘Orus’and autumn -fruit type‘Summit’as experimental materials, with paraffin cut section method, we observed the developing process of flower bud differentiation, development of microspore and megaspore, development of male and female gametophytes, and anatomical structure of fruit. The changes of stored pollen energy under room and low temperature were tested with TTC. Pollen morphology was observed with scanning electron microscope (SEM). Effects of stigma receptivity, effective pollination period, pollination ways on setting fruit rate and seed germination ability under different treatment were studied. The main conclusions as follows:
1. The flower bud differentiation process of Raspberry included pre-differentiation stage, early differentiation and small flower differentiation stage. The small flower differentiation stage included stage of sepal primordium differentiation, stage of stamen primordium differentiation, stage of pistil primordium differentiation made of a lot of apocarpous pistil, and stage of petal primordium differentiation. In Harbin, There was no much difference in the process of flower bud differentiation between 2-years and 1-year cane of‘Summit’and 2-years cane of‘Orus’, except the period of flower bud differentiation.
2. The characteristics of micro -megaspore and male -famale gametophytes: the anther wall was dicotyledonous type and the tapetum was secretory. There were many microspore mother cells, the meiosis tetrads were tetrahedral, microspore was circle and smaller, and the male gametophytes were 2-cell types.The embryo sac was polygonum type , there were two ovules in each ovary, one was degenerate, the other was developped normally, the ovule was anatropous.
3. The mature pollen was circle with three germinal furrows. The exterior wall of pollen was a reticulation. There was a lid on the germination aperture ,and there were small grains on the surface layer of germination furrows with cover.
4. The pollen vitality was up to 50% in‘Summit’whereas decreased to 38.26% and 44.59% in‘Orus’afer the pollens were stored under room temperature and low temperature in the refrigerator for 20 days with TTC method.
5. The stigma receptivity of‘Orus’could last 24 hours, and stigma receptivity of‘Summit’could last 30 hours. The effective pollination period of‘Orus’was 7 days, and that of‘Summit’was 4 days. The self-fruitful percent,natural-fruitful percent and artificial pollination percent of‘Orus’were 52.73%, 84.73% and 72.46% respectively, and that of‘Summit’were 46.33%、90.62%、79.61% respectively.
6. Fruit of Raspberry was composed of pericarp and seed. The pericarp had 3 layers from outside to inside. Exocarp was thin and velvet. Mesocarp was composed of multi-layer parenchyma cell .Endocarp composed of a lot of fibrous scleroid cell . Fruit type was aggregate stone fruit. There was only one seed that has seed coat and embryo. The seed coat was thick and arranged tightly with multi-layer cell.
7. The fruit growth curve of Raspberry was“double S”curve. There was one slow development period between the two fast development periods, but it lasted shortly.
1.树莓花芽形态分化过程可分为花芽未分化期、花芽分化初期、小花分化期三个时期,小花分化期又包括花萼原基、雄蕊原基和大量离生心皮组成的雌蕊原基时期,花瓣原基。在哈尔滨地区,品种撒米特的2年生枝条、1年生枝条和奥瑞斯的2年生枝条花芽分化进程基本一致,分化时间不同。
2.大小孢子发育和雌雄配子体的发育特点为花药壁属于双子叶型,绒毡层为腺质(分泌)绒毡层,小孢子母细胞为多数,减数分裂四分体呈四面体型,小孢子圆形,较小,雄配子体为2-细胞型;蓼型胚囊,每子房内有2个胚珠,其中1个退化,另1个正常发育。倒生胚珠。
3.成熟花粉粒圆型,具有三个萌发沟,花粉外壁呈网状,萌发孔具有孔盖,萌发沟、表层具有小颗粒。
4.采用TTC测定保存在常温下和冰箱保鲜层中的花粉,表明无论在常温下,还是在冰箱保鲜层中保存20天后撒米特的花粉活性仍可达到50%以上,而奥瑞斯的花粉分别降低到38.26%和44.59%。
5.奥瑞斯柱头可授性可持续24小时,撒米特柱头可授性可持续30小时。奥瑞斯有效授粉期为7天,撒米特有效授粉期为4天。奥瑞斯自交结实率、自然结实率和人工授粉结实率分别为:52.73%、84.73%、72.46%;撒米特自交结实率、自然结实率和人工授粉结实率分别为:46.33%、90.62%、79.61%。
6.解剖观察表明树莓果实由果皮和种子两部分组成。果皮自外向内具有三层,外果皮较薄,表面光滑。中果皮由多层薄壁细胞组成,内果皮坚硬,由大量纤维状厚壁细胞组成,果实属于聚合核果类型;种子1枚,由种皮和胚组成,种皮较厚,多层细胞组成,排列紧密。
7.树莓果实的生长曲线表现为双S曲线,在两次速长期之间有1个缓慢生长期,但持续时间短。
Raspberry, small fruit of Rosaceae, Rubus L., has two types according to the time of fruit forming: summer bearing and primocane bearing. Both 2-years and 1-year cane of primocane bearing type can produce fruits and we can use 1-year cane, which need not be buried in winter to reduce the cost. Now, there are some reports about the flower bud differentiation of summer-fruit type Raspberry in China and there is still a blank in the study of Raspberry procreate biology. In this paper, we took summer-fruit type‘Orus’and autumn -fruit type‘Summit’as experimental materials, with paraffin cut section method, we observed the developing process of flower bud differentiation, development of microspore and megaspore, development of male and female gametophytes, and anatomical structure of fruit. The changes of stored pollen energy under room and low temperature were tested with TTC. Pollen morphology was observed with scanning electron microscope (SEM). Effects of stigma receptivity, effective pollination period, pollination ways on setting fruit rate and seed germination ability under different treatment were studied. The main conclusions as follows:
1. The flower bud differentiation process of Raspberry included pre-differentiation stage, early differentiation and small flower differentiation stage. The small flower differentiation stage included stage of sepal primordium differentiation, stage of stamen primordium differentiation, stage of pistil primordium differentiation made of a lot of apocarpous pistil, and stage of petal primordium differentiation. In Harbin, There was no much difference in the process of flower bud differentiation between 2-years and 1-year cane of‘Summit’and 2-years cane of‘Orus’, except the period of flower bud differentiation.
2. The characteristics of micro -megaspore and male -famale gametophytes: the anther wall was dicotyledonous type and the tapetum was secretory. There were many microspore mother cells, the meiosis tetrads were tetrahedral, microspore was circle and smaller, and the male gametophytes were 2-cell types.The embryo sac was polygonum type , there were two ovules in each ovary, one was degenerate, the other was developped normally, the ovule was anatropous.
3. The mature pollen was circle with three germinal furrows. The exterior wall of pollen was a reticulation. There was a lid on the germination aperture ,and there were small grains on the surface layer of germination furrows with cover.
4. The pollen vitality was up to 50% in‘Summit’whereas decreased to 38.26% and 44.59% in‘Orus’afer the pollens were stored under room temperature and low temperature in the refrigerator for 20 days with TTC method.
5. The stigma receptivity of‘Orus’could last 24 hours, and stigma receptivity of‘Summit’could last 30 hours. The effective pollination period of‘Orus’was 7 days, and that of‘Summit’was 4 days. The self-fruitful percent,natural-fruitful percent and artificial pollination percent of‘Orus’were 52.73%, 84.73% and 72.46% respectively, and that of‘Summit’were 46.33%、90.62%、79.61% respectively.
6. Fruit of Raspberry was composed of pericarp and seed. The pericarp had 3 layers from outside to inside. Exocarp was thin and velvet. Mesocarp was composed of multi-layer parenchyma cell .Endocarp composed of a lot of fibrous scleroid cell . Fruit type was aggregate stone fruit. There was only one seed that has seed coat and embryo. The seed coat was thick and arranged tightly with multi-layer cell.
7. The fruit growth curve of Raspberry was“double S”curve. There was one slow development period between the two fast development periods, but it lasted shortly.
引文
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