摘要
普通油茶(Camellia oleifera)是我国特有的木本油料树种,其树体及果实的利用价值在适生区的生态及林业产业建设中具有重要的地位。本文对当前生产中的主要普通油茶无性系开花生物学及可配性进行了研究和分析,研究内容包括无性系的开花物候、花粉的生长特性、花粉的形态分类、花粉的贮藏条件、无性系之间的可配性等方面,以期对花粉的发育和配置提供理论依据,研究得出了以下结果。
普通油茶花期开始于10月下旬,结束于12月中旬个别到1月上旬,花期长达80d,属于“持续开花模式”型物种,有利于传粉和授粉,但授粉过程容易受到气候的影响；普通油茶花果同期,果实一般成熟于10月中旬,不晚于盛花期；相同无性系在不同年份间花期物候有差异。
通过对2009和2010年的花期聚类分析,普通油茶无性系的花期可分为早、中、晚3类。所试无性系花期类型较为固定的为,21号、23号、53号、55号、88号、100号、297号无性系均属于早花种,4号、10号、27号、40号、178号均属于中花种,82号、297号属于晚花种,3号、100号、185号、213号无性系花期类型年份间有不同。
采用正交试验研究了6个普通油茶无性系花粉不同温度、硼元素用量、蔗糖浓度、琼脂用量对花粉萌发的影响。结果表明,温度为所有无性系花粉萌发的主要影响因子,对萌发率的影响最大；蔗糖浓度对除82号以外无性系的花粉萌发率影响显著；硼元素浓度仅对178号无性系的萌发率的影响达极显著水平,琼脂用量仅对185号无性系的萌发率影响显著。根据因素对各参试无性系花粉萌发的影响程度,可以确定4个因素的主次顺序为：温度>蔗糖浓度>硼元素用量~琼脂用量。各因素水平对油茶花粉萌发率的影响效果为：温度25℃>18℃>12℃：蔗糖浓度10%>5%-15%；硼元素以100 mg·kg-1的用量最佳；琼脂用量以0.5%为最佳。结合多重比较,筛选出普通油茶花粉萌发的最适培养组合为T9,即100 mg.kg-1硼元素用量+10%蔗糖浓度+0.5%琼脂用量培养基+25℃恒温培养。
利用普通显微镜和荧光显微镜对普通油茶在最适培养基培养下和授粉后不同时间段花粉管的生长状况进行了观察。结果显示,普通油茶花粉在最适的培养条件下即可生长出花粉管,花粉管的生长速率在不同时间段差异显著,开始时生长略缓,12h左右生长加速,22h后生长越来越趋于平缓,萌发后24h花粉管基本停止生长,整个花粉管的生长过程呈“慢—快—慢”的抛物线趋势图；观察普通油茶花粉授粉后花粉萌发荧光显示：一般授粉后0.5h开始萌发长出花粉管,随后花粉管仅管长不断的伸长,管径的变化不大,约24h穿过花柱基部进入子房室,约48h即能完成受精过程；普通油茶花粉授粉后的花粉管的生长状况呈现为“快—慢”的趋势图,与离体生长有差异,花粉在柱头授粉后的生长量与生长速率与离体萌发差异显著,活体萌发相对于离体培养生长的更快,更准确的反映油茶花粉的生长动态。
以6个普通油茶无性系花粉为试验材料,研究了不同浓度的无机元素(钙、锌、钼)对花粉萌发率的影响,结果表明,钙离子和锌离子浓度对普通油茶花粉的萌发表现出一定的抑制作用,随着钙离子浓度的增加花粉的萌发率下降；较低的钙、锌离子浓度有利于普通油茶花粉的萌发,高于100 mg·L-1的钼离子也抑制花粉萌发。
本文研究了6个普通油茶无性系花粉在不同贮藏处理条件下花粉生活力随着时间变化的规律。结果表明,贮藏后花粉生活力的下降分为两个阶段,前30d失活快,下降明显；30d以后下降逐渐放慢,最后趋于平缓。整个过程表现为“快—慢—平缓”的趋势。花粉的贮藏因素中,温度对贮藏效果的影响最大,低温有利于花粉活力的保持,湿度对贮藏效果也有影响,具干燥剂的瓶装密封处理利于贮藏；经过对9种贮藏方式的优选,结果为冷冻状态下的瓶装密封硅胶干燥(处理9)方式为最佳贮藏处理,在此贮藏方式下,普通油茶花粉贮藏一年后仍能保持约10%的生活力。
利用扫描电子显微镜对3个普通油茶优良品系的花粉形态进行了观测,结果表明,普通油茶花粉为中型花粉,大小为35.42～43.75μm×36.46～42.71μm；P/E值为0.83～1.11,形状为球形或近球形。赤道面观为椭圆形或卵圆形,极面观三裂圆形或近圆形。具3条沿极轴方向走行、等间距作120°等份分布的孔沟,长度几达花粉两极,宽6.30～11.6μm；沟中可见细密的颗粒,无沟膜或沟膜不明显,属于N3P4C3型花粉。外壁纹饰为脑纹或桔皮状,3种普通油茶无性系花粉之间形状和外壁纹饰有显著差异。
通过对普通油茶主要无性系的自然座果率的统计观察,结果显示,普通油茶的自然座果普遍不高,着果率为16.33%,成果率为13.84%。不同无性系之间座果差异显著,无性系40号、185号、53号和4号表现较好,其余无性系表现较差,均低于平均值,178号、185号的落果率较低。
对普通油茶主要无性系间杂交后的调查结果表明,普通油茶杂交对座果提高不明显,相对于母本着果率平均增加4.41%,成果率仅提高1.90%：相对于总体着果率平均增加3.57%,成果率仅提高1.23%。个别杂交组合表现较好,185号作为母本与81号杂交组合的座果效果最好,着果率达60.44%,成果率达55.11%；其次为53号作为母本分别于185号和81号杂交组合次之；以40号和82号做母本的杂交效果比母本的自然座果效果差,需避免。结果还显示,通过杂交并不能明显的降低落果率。
以普通油茶无性系为母本与小果油茶杂交均可座果,二者应属近缘种。杂交组合中以4号无性系为母本与小果油茶的杂交组合座果效果较好,其中4×M9着果率达30%,成果率为22.50%,座果效果最好；以82号无性系作为母本的杂交组合座果的效果次之。小果油茶与普通油茶的杂交效果相对于普通油茶自然座果效果要差,杂交后更易落果。
C.oleifera oleifera is a characteristic arboreal oil tree plant of China, Its tree body and the using value of the fruit plays an important role in living area and the construction in the forestry industry. This paper directed to the biological character of blossom and the crossability test of C.oleifera, mainly researched the flowering phenology of the main clones, the characteristics of pollens germination, the morphology of pollens, the pollens storage conditions, the Crossability of main clones, except to provide the theory in the pollens growth and in hybrid. The results were as follows.
The flowering duration of populations spanned from late October to late mid-December, the flowering phenology was 80d individually, belonged "Mass-flowering" species. it was advantageous in the pollination and the pollination, but the pollination process easy to receive the climate the influence; the C.oleifera's blossom and fruit were in the same period, fruit general matured in mid-October, but not late in peak flowering phenology; the flowering phenology of same clones has differences between 2009 an 2010.
Through to 2009 and 2010 flowering season cluster analysis, the C.oleifera may divide into the morning, and evening 3 kinds. The type of tries the clone flowering phenology was fixed,21,23,53, 55,88,100,297 clones belonged to the early flowering phenology,4,10,27,40,178 clones belonged to the noon flowering phenology,82,297 clones belonged to the late flowering phenology, 3,100,185,213 clones flowering phenology had the difference between 2009 an 2010.
To study the characteristics of pollen germination in vitro of 6 C.oleifera species, the orthogonal test using 6 C. oleifera species as the materials was aimed to examine the effects of the 4 factors consisting of temperature, boron consumption, sucrose, and agar usage on pollen germination. The results indicated that temperature was the most significant factor of all species among the 4 factors, sucrose markedly affected the pollen germination of most species except 82 clones, and boron consumption and agar usage markedly impacted on 178 clones and 185 clones respectively.Comparing to the 4 factors impact on each test clones, the primary sequence of each factors, temperature was 25℃>18℃>12℃; sucrose was 10%>5%- 15%; the optimum of boron consumption and agar usage were 100 mg·kg-1 and 0.5%. Combining multiple comparision, the optimum extractive condition is T9:100 mg·kg-1 boron consumption +10% sucrose +0.5% agar usage+25℃temperature.
To study the micro dynamic pollination and fertilization of the C.oleifera Abel, the growth of pollen on stigma were observed by optical microscope and electron microscope. The results indicated that the pollen tube percentage was significant different in different period, at the beginning the growth was slow, about the 12 hours accelerate the growth,after 22 hours the growth trend was gentle, after 24 hours the growth stopped. The whole growth of pollen tube followed the "slow-fast-slow" trent. The pollens began to germinate 0.5 hour after pollination and the pollen tubes penetrated half of the style 8 hours after pollination and reached the style base, entered the ovary after 24 hours, and the fertilization was completed about 48 hours after pollination. The whole growth of pollen tube followed the "fast-slow" trent. The germination and growth of pollen on stigma was faster significantly than that of in vitro, it could more accurately react the actual growth situations.
To study on the effects of calcium (Ca2+)、zinc(Zn2+)and molybdenum (Mo6+)on pollen germination percentage of 6 C.oleifera species. The results indicated that when the concentration of Ca2+ was high, the pollen germination was centration inhibited, If the concentration of Zn2+、Mo6+ was low, the pollen tube growth would be enhanced. But when the concentration of Zn2+、Mo6+ was high, the pollen germination of pollen tube was inhibited.
This paper analyzed pollen viability of 6 ordinary clone C.oleifera in different storage condition as times changed. The results showed that pollen viability can be divided into 2 stages, the viability declined fast in the first 30d, then declined slower, and ended in stable. The whole process was as fast to slow to stable. In the storage, temperature was the key element, low temperature was good for viability. Moisture also affected the viability, it's good to seal the pollen in the bottle with feature of dry. The best storage condition was to seal the pollen in silica-gel drier bottle in frozen, in which the ordinary clone C.oleifera pollen could have 10% viability after 1 year.
To study on the evolution status of C.oleifera in Theacea, pollens of 3 C.oleifera species were scaned by electron microscope. The results showed that the shape of pollen was mainly prolate to subsphemidal from the equatorial view, the pollen grain size were middle, sizes was 35.42～43.75μm×36.46～42.71μm;P/E was 0.83～1.11. The equatorial view was oval or ovoid, polar view was tri-lobed circular or circular, the aperture was 3-colporat containing granulats which directed along the polar axis of 120°almost stretching 2 poles, which length was 6.30～11.671μm, the colporat contained some granulats which had not the membrane or not obvious, the shape was N3P4C3. The orange surface or brain veins sculpture which indicated that C.oleifera might be primitive groups in the Theacea. There existed significant diferences in the shape and the ornamentation.
Based on the natural average fruit-set rate statistical observation of main ordinary clone C.oleifera, the results showed that the average C.oleifera natural fruit-set generally was not high, fruit-bearing rate was 16.33%, and fruit-receiving rate 13.84%. Difference among clones was significant. Fruit-set clone no.40,185,53 and 4 are good, the rest even lower than the average, no. 178 and 185 full-drop ratio were lower.
According to investigation of C.oleifera hybridization between the main clone, the results showed that C.oleifera of fruit-set improve cross incompatibilities was not obvious, compared to the maternal rate, the average fruit-bearing rate increased 4.41%, fruit-receiving rate only 1.90%; compared to the overall rate, the average fruit-bearing rate increased 3.57% fruit-receiving rate 1.23% only. Some specific hybrid combinations were better, the fruit-set effect of cross combinations of 185 and 81 was the best, fruit-bearing rate was 60.44% and fruit-receiving rate was 55.11%, the next is hybrid combinations of no.53 as maternal respectively with 185 and 81; the hybrid combinations of 40 and 82 was poor, need to be avoided. The results revealed that, hybridization can't significantly reduce full-drop rate.
In C.oleifera clone for maternal and C.meiocarpa hybrid all could fruit-set, both of allied should belong to. fruit-set effect of hybrid combinations of 4 clone and C.meiocarpa as maternal was good, where the fruit-bearing rate of 4 x M9 was up to 30%, fruit-receiving rate 22.50%, the best fruit-set effect. The next was fruit-set effect of Hybrid combinations of 82 as one of the maternal. Compared to conventional hybrid effect of natural fruit-set effects, C.meiocarpa with C.oleifera was poorer, crossbred fluctuation easier.
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