黑果枸杞两种花型的花部综合征与传粉特性

doi:10.17521/cjpe.2021.0463

植物生态学报 ›› 2022, Vol. 46 ›› Issue (9): 1050-1063.DOI: 10.17521/cjpe.2021.0463

黑果枸杞两种花型的花部综合征与传粉特性

哈里布努尔¹, 古丽扎尔·阿不都克力木¹^,², 热依拉穆·麦麦提吐尔逊¹, 艾沙江·阿不都沙拉木¹^,²^,^*()

¹喀什大学生命与地理科学学院, 新疆喀什 844000
²新疆帕米尔高原生物资源与生态重点实验室, 新疆喀什 844000

收稿日期:2021-12-09 接受日期:2022-05-07 出版日期:2022-09-20 发布日期:2022-10-19
通讯作者: 艾沙江·阿不都沙拉木
作者简介:* (aysajanxj@sina.com)
基金资助:
国家自然科学基金(31860121);“天山雪松计划”青年拔尖人才后备人选培养项目和新疆维吾尔自治区高层次人才培养计划项目(QN2016BS0597)

Flower syndrome and pollination characteristics of two flower morphs in Lycium ruthenicum (Solanaceae)

Halibunuer ¹, Gulzar ABDUKIRIM¹^,², Reyilamu MAIMAITITUERXUN¹, Aysajan ABDUSALAM¹^,²^,^*()

¹College of Life and Geographic Sciences, Kashi University, Kashi, Xinjiang 844000, China
²Key Laboratory of Biological Resources and Ecology of Pamirs Plateau in Xinjiang Uygur Autonomous Region, Kashi, Xinjiang 844000, China

Received:2021-12-09 Accepted:2022-05-07 Online:2022-09-20 Published:2022-10-19
Contact: Aysajan ABDUSALAM
Supported by:
National Natural Science Foundation of China(31860121);“Tianshan Xuesong” Young Top Talent Reserve Project and the High-level Personnel Training Program of the Xinjiang Uygur Autonomous Region(QN2016BS0597)

摘要/Abstract

摘要：

黑果枸杞(Lycium ruthenicum)是中国西北地区极端环境中分布的国家二级保护植物, 该物种在新疆南部的自然种群中出现了同型花柱类型(同位花)和柱头探出式雌雄异位类型(异位花)个体, 并且遭遇沙尘暴频繁的种群中异位花个体出现频率减少。该研究对喀什市自然种群中黑果枸杞两种不同花型植株的花部综合征和传粉特性进行比较研究, 以期探讨该物种不同花型植株在南疆早春极端环境中的花部特征的可塑性及其适应性机制。结果表明: 同位花雌雄蕊高度间无显著差异, 而异位花雌蕊高度显著高于雄蕊; 同位花花冠直径、花冠筒长、胚珠数均高于异位花, 而异位花雌雄蕊空间距离、花粉数及花粉胚珠比均比同位花高。黑果枸杞同位花个体比例(68%)高于异位花个体(32%), 种群水平及个体水平同位花花期((117.00 ± 2.25) d, (101.65 ± 1.98) d)比异位花((26.00 ± 1.00) d, (18.75 ± 1.00) d)长, 而单花水平上异位花单花寿命((4.50 ± 0.14) d)比同位花((3.13 ± 0.11) d)长。两种类型花在花早期(紫色)分泌的花蜜量均高于花后期(白色)。在紫色花阶段(花开放早期), 同位花上的主要传粉者意大利蜜蜂、熊蜂和食蚜蝇的访花频率和停留时间均高于异位花; 而白色花阶段(花开放后期)意大利蜜蜂、熊蜂在异位花上的访花频率比同位花高。在不同花色阶段, 同位花柱头花粉落置数、花粉移出率、花粉传递效率均比异位花高, 并且同位花自然坐果率及结籽率均比异位花高。在新疆南部的沙尘暴极端环境下, 同位花通过较高的自交亲和性保障繁殖, 而异交为主的异位花提高了异交率。异位花与同位花在花部综合征和花报酬上的差异, 是影响其繁殖成功的主要因素。

关键词: 黑果枸杞, 同位花柱, 柱头探出式雌雄异位, 传粉特性

Abstract:

Aims Flower syndrome and pollination characteristics are the basis and main driving forces of the evolutionary success of flowering plants. Lycium ruthenicum is a national second-class protected plant distributed in northwest China, with two flower morphs, i.e. homostylous flower (style length is similar to stamen length) and flower with approach herkogamy (style length is longer than stamen length). However, little is known about the flower syndrome and pollination characteristics of the different flower types of L. ruthenicum.

Methods In this study, we compared the flowering phenology, flower characteristics, flower reward, and pollination characteristics of two different flower morphs of L. ruthenicum.

Important findings In Kashi natural populations, there is no significant difference in the height of sexual organs of homostylous flower, but it is significant in the herkogamous flowers. The corolla diameter, corolla tube length, number of ovules of homostylous flowers are higher than that of herkogamous flowers. The stamen-stigma distance, number of pollen and pollen/ovule (P/O) of herkogamous flowers are higher than that of homostylous flower. The proportion of homostylous individuals (68%) is higher than that of herkogamous (32%). Population and individual level flowering times were significantly different between the flower morphs and were longer for homostylous flower individuals ((117.00 ± 2.25) d, (101.65 ± 1.98) d) than that of herkogamous flower individuals ((26.00 ± 1.00) d, (18.75 ± 1.00) d), but the longevity of herkogamy flowers ((4.50 ± 0.14) d) are longer than that of homostylous flowers ((3.13 ± 0.11) d). At early flower stage (purple phase) the nectar volume of two morphs is significantly higher than late stage (white phase). In the purple phase the main pollinators were Bombussp., Apis mellifera and Syrphidae, and the visiting frequency and duration of stay on the homostylous flowers are higher than that of herkogamy flowers. In the white phase, the main pollinators on the herkogamous flower have higher visiting frequency than that of homostylous flower. At the different phases the number of pollen deposited on the stigma, pollen removal rate, pollination efficiency of homostylous flowers are higher than that of herkogamous flowers. The fruit and seed set of homostylous flowers are higher than that of herkogamous flowers. Under the extreme environment of sandstorms in southern Xinjiang, homostylous flowers can secure reproduction through high self-compatibility, and herkogamous flowers will increase outcrossing rate in the population. These differences in flower traits and pollination adaptations may be the main factor affecting the reproductive success of L. ruthenicum.

Key words: Lycium ruthenicum, homostylous, approach herkogamy, pollination characteristic

哈里布努尔, 古丽扎尔·阿不都克力木, 热依拉穆·麦麦提吐尔逊, 艾沙江·阿不都沙拉木. 黑果枸杞两种花型的花部综合征与传粉特性. 植物生态学报, 2022, 46(9): 1050-1063. DOI: 10.17521/cjpe.2021.0463

Halibunuer , Gulzar ABDUKIRIM, Reyilamu MAIMAITITUERXUN, Aysajan ABDUSALAM. Flower syndrome and pollination characteristics of two flower morphs in Lycium ruthenicum (Solanaceae). Chinese Journal of Plant Ecology, 2022, 46(9): 1050-1063. DOI: 10.17521/cjpe.2021.0463

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0463

https://www.plant-ecology.com/CN/Y2022/V46/I9/1050

图/表 9

图1 黑果枸杞花各部位特征测量(A)和不同花型花花柱及花药相对高度(B)(平均值±标准误)。a(a′)-e(e′), 花冠筒长度; a(a′)-d(d′), 雌蕊长; a(a′)-c(c′), 雄蕊长; f(f′)-b(b′), 花冠口直径。B中不同小写字母表示同一花型花不同性器官间差异显著(p < 0.05)。

Fig. 1 Measurements of flower traits (A) and relative height of sexual organs (B) of two flower morphs of Lycium ruthenicum (mean ± SE). a(a′)-e(e′), flower length; a(a′)-d(d′), pistil length; a(a′)-c(c′), stamen length; f(f′)-b(b′), corolla diameter. Different lowercase letters in B indicate significant difference between different sexual organs of the same flower morph (p < 0.05).

图2 新疆喀什黑果枸杞的植株、花序、果实、花型及花颜色变化。A, 雌雄异位花。B, 同位花。C, 异位花花序。D, 同位花花序。E, F, 果实。G, 植株。H, 不同花型花在花寿命内的花色变化(上部为同位花, 下部为异位花)。

Fig. 2 Individual plant, inflorescence, fruit, flower morphs and flower color changes in Lycium ruthenicum in Kashi, Xinjiang, China. A, Herkogamous flower. B, Homostylous flower. C, Inflorescence of herkogamous flower. D, Inflorescence of homostylous flower. E, F, Fruit. G, Individual plant. H, Change of flower color in different phases of flower longevity in two flower morphs of L. ruthenicum (upper, homostylous flower; lower, herkogamous flower).

表1 黑果枸杞同位花和雌雄异位花在花寿命早期(紫色)和后期(白色)花部特征(平均值±标准误)的比较(广义线性模型)

Table 1 Comparisons of flower traits (mean ± SE) among two flower morphs at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum using generalized linear model (GLM)

花部特征 Flower characteristic	紫色阶段 Purple phase				白色阶段 White phase				花型×阶段 Morph × phase
花部特征 Flower characteristic	同位花 Homostylous flower	雌雄异位花 Herkogamous flower	Wald χ₁²	p₁	同位花 Homostylous flower	雌雄异位花 Herkogamous flower	Wald χ₂²	p₂	Wald χ₃²	p₃
花冠直径 Corolla diameter (mm)	8.29 ± 0.25	7.36 ± 0.33	10.411	0.001	7.18 ± 0.23	6.70 ± 0.43	0.914	0.659	17.380	0.001
花冠筒长 Corolla tube length (mm)	14.28 ± 0.31	10.70 ± 0.54	32.514	<0.001	14.92 ± 0.39	9.71 ± 0.57	57.284	<0.001	4.188	0.242
雌蕊长 Pistil length (mm)	11.99 ± 0.33	12.20 ± 0.57	0.104	0.747	12.06 ± 0.33	13.07 ± 0.48	3.005	0.083	0.043	0.998
雄蕊长 Stamen length (mm)	11.99 ± 0.24	10.20 ± 0.42	13.574	<0.001	12.37 ± 0.36	11.14 ± 0.53	3.745	0.053	1.481	0.687
雌雄蕊空间距离 Stamen-stigma distance (mm)	0.65 ± 0.16	2.00 ± 0.27	18.588	<0.001	1.02 ± 0.18	1.93 ± 0.27	8.042	0.005	4.358	0.225
雌蕊-花冠口距离 Corolla tube-stigma distance (mm)	2.29 ± 0.22	1.50 ± 0.39	3.175	0.075	2.86 ± 0.28	3.36 ± 0.42	0.975	0.323	12.430	0.006
雄蕊-花冠口距离 Corolla tube-stamen distance (mm)	0.70 ± 0.28	2.29 ± 0.16	24.444	<0.001	2.55 ± 0.29	1.86 ± 0.42	1.865	0.172	1.208	0.751

图3 黑果枸杞不同花型花紫色阶段(1、2和3)和白色阶段(4和5)花粉活力及柱头可授性比较(平均值±标准误)。“+”表示柱头可授性强弱程度。

Fig. 3 Pollen viability and stigma acceptability in purple phase (1, 2 and 3) and white phase (4 and 5) of two flower morphs of Lycium ruthenicum (mean ± SE). “+” indicates the degree of acceptability of stigma.

图4 黑果枸杞不同花型花在花寿命早期(紫色)和后期(白色)的花蜜分泌量(平均值±标准误)。不同大写字母表示同种类型不同阶段间的差异显著, 不同小写字母表示同一阶段不同花型间的差异显著(p < 0.05)。

Fig. 4 Nectar volume at early flowering phase (purple) and late phase (white) of two flower morphs of Lycium ruthenicum (mean ± SE). Different uppercase letters indicate significant difference between different phases of the same flower morph, and different lowercase letters indicate significant difference between different flower morphs at the same phase (p < 0.05).

表2 新疆喀什黑果枸杞不同花型花在花寿命不同阶段(紫色和白色)花气味中的化学成分占总化合物百分比(平均值±标准误)的比较

Table 2 Mean relative composition of two flower morphs at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum flower volatiles in Kashi, Xinjiang, China (mean ± SE)

传感器 Sensor	物质 Substance	紫色阶段 Purple phase				白色阶段 White phase				花型×阶段 Morph × phase
传感器 Sensor	物质 Substance	同位花 Homostylous flower	雌雄异位花 Herkogamy flower	Wald χ₁²	p₁	同位花 Homostylous flower	雌雄异位花 Herkogamy flower	Wald χ₂²	p₂	Wald χ₃²	p₃
W1C	芳香成分苯类 Benzenoids	8.58 ± 0.37	9.26 ± 0.46	1.408	0.235	6.31 ± 0.20	8.23 ± 0.04	130.437	<0.001	3.385	0.066
W3C	氨类, 芳香成分 Ammonia, benzenoids	8.19 ± 0.40	7.01 ± 1.49	0.463	0.496	5.45 ± 0.27	7.76 ± 0.05	106.682	<0.001	3.120	0.077
W5C	烷烃芳香成分 Alkane aromatic composition	8.06 ± 0.41	8.92 ± 0.55	1.566	0.211	5.28 ± 0.28	7.64 ± 0.05	103.141	<0.001	3.424	0.064
W2W	芳香成分, 有机硫化物 Benzenoids, organic sulfide	10.92 ± 0.39	10.11 ± 0.27	4.025	0.045	14.97 ± 0.24	10.28 ± 0.22	309.736	<0.001	87.539	<0.001
W1S	甲基类 Methyl	19.83 ± 0.55	15.87 ± 1.60	4.450	0.035	25.47 ± 1.01	19.86 ± 0.31	42.106	<0.001	0.531	0.466
W1W	无机硫化物 Inorganic sulfide	7.61 ± 2.61	10.25 ± 0.22	2.484	0.115	13.67 ± 0.19	10.48 ± 0.14	267.990	<0.001	9.389	0.002
W2S	醇类, 醛酮类 Alcohols, aldehydes and ketones	10.40 ± 0.30	10.67 ± 0.14	1.223	0.269	10.92 ± 0.39	10.70 ± 0.10	0.357	0.550	0.279	0.598
W3S	烷烃 Alkane	8.28 ± 0.40	8.99 ± 0.53	1.131	0.288	5.37 ± 0.27	7.77 ± 0.04	112.877	<0.001	4.695	0.030
W5S	氮氧化合物 Nitrogen oxides	9.68 ± 0.29	9.85 ± 0.33	0.160	0.689	7.41 ± 0.36	9.35 ± 0.09	40.361	<0.001	10.389	0.001
W6S	氢化物 Hydride	8.46 ± 0.39	9.08 ± 0.51	0.952	0.329	5.46 ± 0.26	7.93 ± 0.05	125.947	<0.001	6.120	0.013

表2 新疆喀什黑果枸杞不同花型花在花寿命不同阶段(紫色和白色)花气味中的化学成分占总化合物百分比(平均值±标准误)的比较

传感器 Sensor	物质 Substance	紫色阶段 Purple phase				白色阶段 White phase				花型×阶段 Morph × phase
传感器 Sensor	物质 Substance	同位花 Homostylous flower	雌雄异位花 Herkogamy flower	Wald χ₁²	p₁	同位花 Homostylous flower	雌雄异位花 Herkogamy flower	Wald χ₂²	p₂	Wald χ₃²	p₃
W1C	芳香成分苯类 Benzenoids	8.58 ± 0.37	9.26 ± 0.46	1.408	0.235	6.31 ± 0.20	8.23 ± 0.04	130.437	<0.001	3.385	0.066
W3C	氨类, 芳香成分 Ammonia, benzenoids	8.19 ± 0.40	7.01 ± 1.49	0.463	0.496	5.45 ± 0.27	7.76 ± 0.05	106.682	<0.001	3.120	0.077
W5C	烷烃芳香成分 Alkane aromatic composition	8.06 ± 0.41	8.92 ± 0.55	1.566	0.211	5.28 ± 0.28	7.64 ± 0.05	103.141	<0.001	3.424	0.064
W2W	芳香成分, 有机硫化物 Benzenoids, organic sulfide	10.92 ± 0.39	10.11 ± 0.27	4.025	0.045	14.97 ± 0.24	10.28 ± 0.22	309.736	<0.001	87.539	<0.001
W1S	甲基类 Methyl	19.83 ± 0.55	15.87 ± 1.60	4.450	0.035	25.47 ± 1.01	19.86 ± 0.31	42.106	<0.001	0.531	0.466
W1W	无机硫化物 Inorganic sulfide	7.61 ± 2.61	10.25 ± 0.22	2.484	0.115	13.67 ± 0.19	10.48 ± 0.14	267.990	<0.001	9.389	0.002
W2S	醇类, 醛酮类 Alcohols, aldehydes and ketones	10.40 ± 0.30	10.67 ± 0.14	1.223	0.269	10.92 ± 0.39	10.70 ± 0.10	0.357	0.550	0.279	0.598
W3S	烷烃 Alkane	8.28 ± 0.40	8.99 ± 0.53	1.131	0.288	5.37 ± 0.27	7.77 ± 0.04	112.877	<0.001	4.695	0.030
W5S	氮氧化合物 Nitrogen oxides	9.68 ± 0.29	9.85 ± 0.33	0.160	0.689	7.41 ± 0.36	9.35 ± 0.09	40.361	<0.001	10.389	0.001
W6S	氢化物 Hydride	8.46 ± 0.39	9.08 ± 0.51	0.952	0.329	5.46 ± 0.26	7.93 ± 0.05	125.947	<0.001	6.120	0.013

图5 黑果枸杞主要访花昆虫。A, 意大利蜜蜂。B, 熊蜂。C, 食蚜蝇。D, 淡脉隧蜂。E, 丝光绿蝇。

Fig. 5 Main visiting insects of Lycium ruthenicum. A, Apis mellifera. B, Bombus sp. C, Syrphidae. D, Lasioglossum occidens. E, Lucilia sericata.

图6 黑果枸杞不同花型花主要传粉者在花寿命早期(紫色)和后期(白色)的访花频率及停留时间(平均值±标准误)。不同大写字母表示同种花型不同传粉者间的差异, 不同小写字母表示同一传粉者不同花型间的差异显著(p < 0.05)。

Fig. 6 Visiting frequency of main pollinators at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum of two flower morphs (mean ± SE). Different uppercase letters indicate significant difference between different pollinators of the same morph, and different lowercase letters indicate significant difference between different flower morphs of same pollinator (p < 0.05).

图7 黑果枸杞不同花型花在花寿命早期(紫色)和晚期(白色)花粉移出率(A)、柱头落置花粉数(B)、花粉传递效率(C)、自然坐果率和结籽率(D)(平均值±标准误)。A-C不同大写字母表示同种花型不同阶段间的差异显著, 不同小写字母表示不同花型间差异显著(p < 0.05)。

Fig. 7 Proportion of total pollen grains removed (A), deposited pollen grains (B), pollen transfer efficiency (C) and fruit and seed set rate (D) in the two flower morphs of Lycium ruthenicum at early flowering phase (purple) and late flowering phase (white)(mean ± SE). Different uppercase letters indicate significant difference between different phases of the same morph, and different lowercase letters indicate significant difference between different flower morph at the same phase (p < 0.05).

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黑果枸杞两种花型的花部综合征与传粉特性

Flower syndrome and pollination characteristics of two flower morphs in Lycium ruthenicum (Solanaceae)

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