热带作物学报

品种Variety	雄蕊数Number of stamen	雄蕊长Length of stamen/mm	雌蕊数Number of pistil	雌蕊长Length of pistil/mm	花粉量（单枚花药）Quantity pollen (one anther)	雌雄等长比例Equal proportion of male and female/%	雌长雄短比例The ratio of long females to short males/%	雌短雄长比例The ratio of shorter females to longer males/%
热研1号	91±8.09	12.72±1.03	3~5	14.90±1.01	1940±174	6	84	10
热研2号	88±3.87	11.76±0.93	3~4	12.32±1.00	3060±220	30	40	30
热研3号	102±6.67	15.06±0.73	3~4	15.12±0.93	2800±151	74	12	14

品种Variety	雄蕊数Number of stamen	雄蕊长Length of stamen/mm	雌蕊数Number of pistil	雌蕊长Length of pistil/mm	花粉量（单枚花药）Quantity pollen (one anther)	雌雄等长比例Equal proportion of male and female/%	雌长雄短比例The ratio of long females to short males/%	雌短雄长比例The ratio of shorter females to longer males/%
热研1号	91±8.09	12.72±1.03	3~5	14.90±1.01	1940±174	6	84	10
热研2号	88±3.87	11.76±0.93	3~4	12.32±1.00	3060±220	30	40	30
热研3号	102±6.67	15.06±0.73	3~4	15.12±0.93	2800±151	74	12	14

指标Index	雄蕊数Number of stamen	雄蕊长Length of stamen	雌蕊长Length of pistil	花粉量Quantity pollen
雄蕊数	1	0.708^**	0.538^**	0.510^**
雄蕊长		1	0.788^**	0.349
雌蕊长			1	-0.174
花粉量				1

指标Index	雄蕊数Number of stamen	雄蕊长Length of stamen	雌蕊长Length of pistil	花粉量Quantity pollen
雄蕊数	1	0.708^**	0.538^**	0.510^**
雄蕊长		1	0.788^**	0.349
雌蕊长			1	-0.174
花粉量				1

开花日期Date of flowering	露白Appear white	开花当天Flowering day	开花后天数Days after flowering
热研1号	3	3	3	3	3	2	2	1	1
热研2号	3	3	3	3	3	2	2	1	1
热研3号	3	3	3	3	3	2	2	1	1

开花日期Date of flowering	露白Appear white	开花当天Flowering day	开花后天数Days after flowering
热研1号	3	3	3	3	3	2	2	1	1
热研2号	3	3	3	3	3	2	2	1	1
热研3号	3	3	3	3	3	2	2	1	1

海南油茶自交与异交的花粉管生长差异分析

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于钊妍 , 龚涵 , 徐玉芬 , 刘艳菊 , 吴坤林 , 曹秋林 , 贾效成 ^*

热带作物学报 | 作物栽培与生理生化 2024,45(12): 2592-2598

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热带作物学报 |作物栽培与生理生化 2024 , 45 (12) : 2592 -2598

海南油茶自交与异交的花粉管生长差异分析

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于钊妍（1995—），女，硕士，研究实习员，研究方向：经济树种生殖发育、种质资源开发利用。

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companyId=1276204756825666100, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国热带农业科学院椰子研究所，海南文昌　571339)])])]

于钊妍, 龚涵, 徐玉芬, 刘艳菊, 吴坤林, 曹秋林, 贾效成^*

作者信息

中国热带农业科学院椰子研究所，海南文昌　571339

通讯作者:

^* 贾效成（JIA Xiaocheng），E-mail：xcjia1@163.com。

Analysis of Pollen Tube Growth Difference Between Self and Crosspollination of Camellia hainanensis

Zhaoyan YU, Han GONG, Yufen XU, Yanju LIU, Kunlin WU, Qiulin CAO, Xiaocheng JIA^*

Affiliations

Coconut Research Institute, Chinese Academy of Tropical Agricultural Sciences, Wenchang, Hainan 571339, China

出版时间: 2024-12-25 doi: 10.3969/j.issn.1000-2561.2024.12.011

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摘要

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油茶（Camellia spp.）是我国重要的木本油料，在保障国家粮油安全中占有重要地位。前人初步研究表明，油茶的自交不亲和性可能是导致油茶结实率低的重要原因之一。本研究拟采用荧光显微镜观察法，结合形态观察，以热研2号、热研3号海南油茶为材料，自交和杂交组合分别为热研3号×热研3号、热研3号×热研2号，进行自交和异交授粉后采样，采用“双载玻片贴合两面观察法”制片后，进行花粉管荧光显微观察，比较二者花粉管生长的差异。结果表明：热研系列海南油茶柱头前端均有不同程度的弯曲，柱头数为3~5个不等，雌蕊长度略大于雄蕊长度；雌蕊长度和雄蕊数量、雄蕊长度呈显著正相关。其单花开放持续5~8 d，花粉散粉时间和柱头具可授性的时间重叠。花粉管荧光观察表明，热研3号海南油茶自交与异交花粉管前期生长速度大致相同，二者生长速度均在授粉后12~24 h内达到顶峰，自36 h后，杂交花粉管生长长度超过自交花粉管，直至生长至子房内。授粉48 h后，自交与异交花粉管均能到达花柱基部，此时已观察到异交花粉管能进入子房，但自交花粉管生长速度则变慢，直至在花柱基部接近子房处生长停滞。该研究结果为海南油茶品种配置和高效栽培提供理论基础。

关键词

海南油茶 / 自交 / 异交 / 花粉管

Abstract

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Camellia spp. is an important woody oil in China, which plays an important role in ensuring national grain and oil security. Previous studies showed that the self-incompatibility of Camellia spp. may be one of the important reasons for the low seed setting rate. In this study, fluorescence microscopy and morphological observation were adopted. C. hainanica Reyan 2 and Reyan 3 were used as the material, and the self-cross and hybrid combinations were Reyan 3×Reyan 3, Reyan 3×Reyan 2, respectively. Samples were collected after self-cross and cross-pollination. The difference of pollen tube growth between the two groups was compared by fluorescence microscopy. The results showed that there were different degrees of curvature at the front end of the stigmas, the number of stigmas ranged from 3 to 5, and the pistil length was slightly larger than the stamen length. The length of pistil was positively correlated with the number and length of stamen. The single flower opening lasted for 5-8 d, and the time of pollen dispersal overlapped with the time of stigma receptivity. Pollen tube fluorescence observation showed that the growth rates of autocross and outcross pollen tubes of Reyan 3 Camellia hainanica were roughly the same in the early stage, and the growth rates of both reached the peak within 12-24 h after pollination. After 36 h, the growth length of hybrid pollen tube exceeded that of autocross pollen tube until it grew into ovary. 48 h after pollination, both inbred and outbred pollen tubes could reach the base of the style, and it was observed that the outbred pollen tubes could enter the ovary, but the growth rate of the inbred pollen tubes slowed down until the growth stopped at the base of the style near the ovary. The results would provide a theoretical basis for the variety configuration and efficient cultivation of C. hainanica.

Key words

Camellia hainanica / self-pollination / cross-pollination / pollen tube

引用本文

于钊妍, 龚涵, 徐玉芬, 刘艳菊, 吴坤林, 曹秋林, 贾效成. 海南油茶自交与异交的花粉管生长差异分析. 热带作物学报, 2024 , 45 (12) : 2592 -2598 . DOI: 10.3969/j.issn.1000-2561.2024.12.011

Zhaoyan YU, Han GONG, Yufen XU, Yanju LIU, Kunlin WU, Qiulin CAO, Xiaocheng JIA. Analysis of Pollen Tube Growth Difference Between Self and Crosspollination of Camellia hainanensis[J]. Chinese Journal of Tropical Crops, 2024 , 45 (12) : 2592 -2598 . DOI: 10.3969/j.issn.1000-2561.2024.12.011

正文

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油茶（Camellia spp.）是山茶属植物中种子含油率较高且有一定栽培面积树种的统称，是我国重要的食用油料树种之一，被誉为“东方油橄榄”，与油棕（Elaeis quineensis）、油橄榄（Olea europaea）、椰子（Cocos nucifera）并称为世界四大木本油料树种。油茶油营养价值高，不饱和脂肪酸含量高达90%以上^[1-2]。海南油茶（Camellia hainanica）是独具热带特色的油茶新种，俗称“山柚”，其茶籽油称为“山柚油”，山柚油口味醇香、品质优良，且已有研究表明其有效成分含量及物质组成等方面显著优于内陆地区广泛种植的普通油茶^[3-5]。

自交不亲和产生的原因多样，从花粉在柱头萌发至受精卵产生的过程中均会发生不育现象，目前经观察确认自交不亲和的植物多为茄科、罂粟科、十字花科和报春花科等，甜菊的自交不亲和可能发生在花粉对柱头的黏附过程、柱头对花粉的识别过程^[6]。欧洲李花粉与雌蕊中的其中一个的S等位基因相同时，导致花粉无法识别或花粉管生长受到抑制^[7]；油茶基因组中的S-RNase基因在自交不亲和后期起到作用，PODs基因可能参与了油茶自交授粉后花粉管程序性死亡^[8-9]。长瓣短柱茶也因为强自交不亲和性导致胚珠败育，推测是当自交花粉落在柱头上时，未接收到信号，使得胚珠内无成熟胚囊，无法完成受精^[10]，该自交不亲和反应发生在花柱基部，表现为合子前自交不亲和性。威宁短柱油茶自交不亲和性是其结实率低的主要原因之一，在杂交育种中应充分考虑这一因素^[11]。

油茶为雌雄同株的异花授粉植物，使得实生苗个体差异较大，性状不稳定，所以生产上多以无性繁殖为主，由此有关油茶的研究主要侧重于丰产栽培技术方面^[12]。但油茶花粉在柱头萌发生长是受精坐果的前提，直接关系到油茶能否结实。因此开展海南油茶自交和异交授粉后花粉管生长行为的研究，对提高结实率具有一定指导意义。目前，国内学者对油茶展开大量研究，明确其具有自交不亲和性，但海南油茶花、果特征和生长结果习性与普通油茶存在明显差异，仍缺乏系统而全面的研究。本研究以海南油茶为研究对象，从形态学的角度对其自交和异交花粉管的生长行为进行荧光显微镜观察，为海南油茶良种配置和杂交育种提供理论基础。

1　材料与方法

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1.1　材料

试验地点位于中国热带农业科学院椰子研究所油茶基地，选择性状良好的优良品种热研3号海南油茶为试材，异花授粉父本品种为热研2号，年初开展抚育工作。其中供试的热研系列油茶为椰子所从海南油茶中选育的优良品系。

1.2　方法

1.2.1　花粉采集

取大蕾期的花朵，剥下花药室内阴干，收集于离心管中，低温干燥保存^[13-14]。

1.2.2　授粉及采样

以热研3号为母本授粉，自交和杂交组合分别为热研3号×热研3号、热研3号×热研2号。盛花期去雄，用毛笔蘸取花粉轻涂在柱头，肉眼可见柱头处有黄色花粉为授粉完成，采集授粉后3、6、12、24、48、72、96、120、144、168 h的雌蕊各5枚，用卡诺固定液固定，固定12~24 h后放入70%乙醇中，4 ℃冰箱保存备用^[15-17]。

1.2.3　荧光显微观察

参考袁德义等^[18]的方法，将油茶花柱经各级酒精复水（50%酒精→30%酒精→蒸馏水），在8 mol/L的NaOH中浸泡2 h软化，蒸馏水洗净后在0.2%的水溶性苯胺蓝染液中染色6 h以上，制作临时切片，在荧光显微镜下观察。

2　结果与分析

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2.1　海南油茶花器官结构

柱头数多为3~4个，少数为5个，柱头前端弯曲，被雄蕊包裹（图1）。不同品种花柱长度、柱头弯曲程度等均有差异，热研2号雌蕊最短，热研3号最长。热研3号油茶雄蕊数量为102（表1），在3个品种中最多，雄蕊长度也最长，为15.06 mm，虽单枚花药花粉量低于热研2号，但在3个品种中，单花花粉量最多，花形也最大。供试的3个品种雌雄蕊相对长度多为雌长雄短或雌雄等长，热研1号约84%的花为雌长雄短，热研3号雌雄蕊空间位置最接近，约74%的花为雌雄等长。

相关性分析结果表明，雄蕊长度和雌蕊长度呈显著正相关，相关性系数为0.788（表2）；雄蕊数量和雄蕊、雌蕊长度均呈显著正相关。因此，柱头越长的海南油茶品系，雄蕊数量也越多，雄蕊长度也越长，一定程度上降低了自交的成功率。此外，热研3号海南油茶的柱头弯曲程度也最小，这是否由于雄、雌蕊之间的空间位置决定的，还有待进一步研究。

2.2　海南油茶花期物候和柱头可授性

海南油茶单花开放从花朵露白至花瓣脱落、雄蕊掉落为止经历约5~8 d（图2）。但油茶群体花期和单花花期长短均与天气有很大关系，温度较高开花提前，温度较低开花则推迟；海南油茶单花遇高温时会加快枯萎，大风雨水天则难以开放。

2022、2023年对试验地的热研系列油茶进行花期物候观察记录，热研1号11月中旬开始开放，11月末达到盛花期；热研2号11月上旬开始开放，11月中旬盛花期，花期较热研1号长，约40 d；热研3号11月中旬开始开放，11月下旬开始盛花期；热研2号和热研3号2个品种间花期重叠时间较长。

观察表明，海南油茶花开放时，花药开裂，花粉散出，至开花第3天时，花粉基本散毕。此时，柱头可授性也最强，从第4天起，柱头可授性开始下降，至第6天开始基本消失（表3）。供试品系的雌、雄蕊成熟时间基本重叠，与前人研究一致，由此推断，海南油茶自交不亲和的主要原因不表现为雌雄异熟。

2.3　自交与异交花粉管生长情况

由试验结果看出，经过水溶性苯胺蓝染色后，自交和异交能够正常萌发的花粉管呈现强烈的蓝色荧光反应。由图3可见，自花授粉后12 h，大量花粉管生长至花柱1/3处；授粉后48 h，花粉管已经生长至花柱基部，之后花粉管生长缓慢或停滞，到达子房上端后，在荧光显微镜下难以观察到。在异花授粉过程中，授粉后12 h，花粉管生长至花柱1/3处（图4），授粉后24 h，有少量花粉管达到花柱1/2处，授粉后48 h，已能观察到部分花粉管穿过花柱基部，进入子房。

研究表明，海南油茶的自交传粉后，未在一开始发生花粉排斥，花粉在柱头萌发和花粉管生长未受到抑制，但最后仍自交败育，可能与其生殖特性相关。

2.4　自交与异交花粉管生长情况比较

热研系列油茶自交和异交花粉管不同时间段的平均生长速度具有明显差异，总体呈现先加快后缓慢的趋势。由图5可见，在24 h之前自交和异交花粉管的生长特性无明显差异，在相同时间内，自交花粉管能达到异交花粉管同样的长度，数量也无明显减少。但在48 h左右，自交与异交花粉管的生长开始产生差异。自交花粉管在24~48 h间生长速度开始减慢，而异交花粉管则顺利到达花柱基部，说明自交花粉管可能受到抑制。

自交和异交在授粉后12~24 h，生长速度达到最大，分别为0.48 mm/h和0.44 mm/h。观察到的自交和异交的花粉管，随着授粉时间延长，花柱尖端肉眼可见有干枯，在荧光显微镜下荧光较弱，但越靠近花柱基部，荧光越强。

3　讨论

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本研究对海南油茶热研花粉萌发和花粉管生长的观察表明，其自交和异交授粉花粉管都能于48 h达到花柱基部，整个过程中平均生长速度相差不大，但不同时间段的花粉管生长速度不一样，自交花粉管在24 h内生长速度快，超过异交花粉管，但随后生长速度下降直至停止。异交花粉管则能一直保持生长，72 h时已生长进子房。此外，花粉管白天生长速度比夜晚快，可能是由于夜晚温度较低且没有光照，由此推断温度和光照也是影响海南油茶花粉管生长的重要因子之一^[19]。

试验中观察到的油茶自交不亲和现象与陈雅等^[20]、高超等^[21-22]的研究结论一致，海南油茶授粉后花粉管于48 h达到花柱基部，这与湘林1号、攸县油茶、华硕系列油茶授粉后花粉管生长速度不一致，推测产生这种差异的原因可能与海南气候、物种特性有关^[23-25]，也可能和植株的自身营养状况有关，使花粉管生长受限导致坐果率低，产生大小年现象。

多数植物的自交不亲和性是由S位点控制，从形态上主要表现为雌雄异熟和雌雄异位。雌雄蕊的长度和雌雄蕊成熟度的差异均能影响植物的自交亲和性^[26-28]。在油茶中，雌、雄蕊的成熟期重叠，且未产生空间上的位置变化，高超等^[21]也发现，油茶自交受限制的情况和遗传学上配子体、孢子体自交不亲和不同，其败育发生在子房处，也就是合子形成以前。在授粉过程中，油茶花粉或花粉管细胞携带自身信息，不断感知来自柱头或花柱道细胞的外部信号，通过不同的信号转导级联反应，顺利完成异花受精过程。已有研究表明，油茶基因组中的S-RNase基因在自交不亲和后期起到作用，PODs基因也可能参与了油茶自交授粉后花粉管程序性死亡，因此油茶自交不亲和过程包含了一个复杂的生理响应和分子调控网络，在这一进程中，还有哪些关键因子参与其中，还有待进一步研究^[29]。但是随着研究的深入，在十字花科和茄科植物中发现，S-RNase的缺失和功能障碍引起花粉S因子的平行突变，使得该物种从自交不亲和转变为自交亲和，在油茶品种中，是否也存在自交亲和的类型^[30]。

除了自交不亲和之外，花期的气候环境条件也影响花粉萌发和花粉管生长行为^[31-32]。油茶的花粉萌发和花粉管的生长受内外部因素的共同调控。连续低温会导致花粉管生长异常，产生严重落花落果现象，坐果率降低。油茶的花粉管萌发和天气关系密切，花粉萌发需要适宜温度，多次授粉结果表明，在授粉前后天气晴朗时，能在花柱内观察到的花粉管数量远多于授粉后降雨的时期，证明天气是授粉中重要因素之一。在自然条件下观察到，各品种坐果率存在一定差异，表明品种间的自交不亲和程度可能存在差别，如能继续深入探究不同品种间的不亲和强度，配合辅助授粉技术，则可进一步明确油茶自交不亲和的作用机理，为海南油茶品种配置和高产栽培提供理论基础。

基金

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海南省自然科学基金青年基金项目(322QN414)
海南省重点研发计划项目(ZDYF2023XDNY054)

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2024年第45卷第12期

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doi: 10.3969/j.issn.1000-2561.2024.12.011

接收时间：2024-07-08
首发时间：2026-06-23
出版时间：2024-12-25

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收稿日期：2024-07-08
修回日期：2024-08-13

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海南省自然科学基金青年基金项目(322QN414)

海南省重点研发计划项目(ZDYF2023XDNY054)

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中国热带农业科学院椰子研究所，海南文昌　571339

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^* 贾效成（JIA Xiaocheng），E-mail：xcjia1@163.com。

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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品种Variety	雄蕊数Number of stamen	雄蕊长Length of stamen/mm	雌蕊数Number of pistil	雌蕊长Length of pistil/mm	花粉量（单枚花药）Quantity pollen (one anther)	雌雄等长比例Equal proportion of male and female/%	雌长雄短比例The ratio of long females to short males/%	雌短雄长比例The ratio of shorter females to longer males/%
热研1号	91±8.09	12.72±1.03	3~5	14.90±1.01	1940±174	6	84	10
热研2号	88±3.87	11.76±0.93	3~4	12.32±1.00	3060±220	30	40	30
热研3号	102±6.67	15.06±0.73	3~4	15.12±0.93	2800±151	74	12	14

品种Variety

雄蕊数Number of stamen

雄蕊长Length of stamen/mm

雌蕊数Number of pistil

雌蕊长Length of pistil/mm

花粉量（单枚花药）Quantity pollen (one anther)

雌雄等长比例Equal proportion of male and female/%

雌长雄短比例The ratio of long females to short males/%

雌短雄长比例The ratio of shorter females to longer males/%

热研1号

91±8.09

12.72±1.03

3~5

14.90±1.01

1940±174

热研2号

88±3.87

11.76±0.93

3~4

12.32±1.00

3060±220

热研3号

102±6.67

15.06±0.73

3~4

15.12±0.93

2800±151

指标Index	雄蕊数Number of stamen	雄蕊长Length of stamen	雌蕊长Length of pistil	花粉量Quantity pollen
雄蕊数	1	0.708^**	0.538^**	0.510^**
雄蕊长		1	0.788^**	0.349
雌蕊长			1	-0.174
花粉量				1

指标Index

雄蕊数Number of stamen

雄蕊长Length of stamen

雌蕊长Length of pistil

花粉量Quantity pollen

雄蕊数

0.708^**

0.538^**

0.510^**

雄蕊长

0.788^**

0.349

雌蕊长

-0.174

花粉量

开花日期Date of flowering	露白Appear white	开花当天Flowering day	开花后天数Days after flowering
热研1号	3	3	3	3	3	2	2	1	1
热研2号	3	3	3	3	3	2	2	1	1
热研3号	3	3	3	3	3	2	2	1	1

开花日期Date of flowering

露白Appear white

开花当天Flowering day

开花后天数Days after flowering

1 d

2 d

3 d

4 d

5 d

6 d

7 d

热研1号

热研2号

热研3号