Floral substances and daytime release rhythms of 3 <i>Chimonanthus</i> species

HU Guiting; YANG Liyuan; REN Guangbing; ZHAO Hongbo

doi:10.11833/j.issn.2095-0756.20240266

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HU Guiting, YANG Liyuan, REN Guangbing, ZHAO Hongbo. Floral substances and daytime release rhythms of 3 Chimonanthus species[J]. Journal of Zhejiang A&F University. doi: 10.11833/j.issn.2095-0756.20240266

Citation:

HU Guiting, YANG Liyuan, REN Guangbing, ZHAO Hongbo. Floral substances and daytime release rhythms of 3 Chimonanthus species[J]. Journal of Zhejiang A&F University. doi: 10.11833/j.issn.2095-0756.20240266

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Floral substances and daytime release rhythms of 3 Chimonanthus species

doi: 10.11833/j.issn.2095-0756.20240266

1.
Zhejiang Provincial Key Laboratory of Germplasm Innovation and Utilization for Garden Plants/Key Laboratory of National Forestry and Grassland Administration on Germplasm Innovation and Utilization for Southern Garden Plants, College of Landscape Architecture, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China
2.
Jinan Park Development Service Center, Jinan 250100, Shandong, China

Received Date: 2024-03-31
Accepted Date: 2024-06-19
Rev Recd Date: 2024-06-19

Abstract

Objective Flower fragrance is an important characteristic of ornamental plants, which is very crucial for plants reproduction. This study aims to reveal the biosynthesis and release mechanism of floral substances in Chimonanthus, and to provide a solid theoretical basis for the utilization and development of Chimonanthus resources. Method Headspace solid phase microextraction (HS-SPME) and gas chromatography-mass spectrometry (GC-MS) were used to identify and analyze floral substances of 3 Chimonanthus species, including C. praecox, C. salicifolius and C. nitens. Result A total of 44 floral substances were identified in 3 Chimonanthus species, among which 19, 23 and 18 substances were identified in C. praecox, C. salicifolius and C. nitens respectively. The floral substances of alcohols and esters were the highest in C. praecox, while the content of alkenes was the highest in the floral substances of C. salicifolius and C. nitens. The daytime release rhythms of floral substances in 3 Chimonanthus species were different. The floral substances of C. salicifolius and C. nitens in the morning, noon and afternoon were different, but the floral substances of C. praecox changed little. Phylogenetic analysis showed that C. salicifolius and C. nitens were more closely related than C. praecox. Conclusion The floral substances of C. salicifolius and C. nitens are similar, which caters to the phylogenetic relationship between them. In view of the special low-temperature flowering environment of C. praecox, the interaction between pollination insects and floral fragrance may be the reason for the uniqueness of floral substances. [Ch, 3 fig. 3 tab. 35 ref.]
- Chimonanthus,
- floral substances,
- release rhythms

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沈阳化工大学材料科学与工程学院沈阳 110142

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蜡梅属Chimonanthus是中国的特有属，公认的蜡梅属植物有4个物种，即蜡梅C. praecox、山蜡梅C. nitens、柳叶蜡梅C. salicifolius和西南蜡梅C. campanulatus^[1−2]。蜡梅广泛分布于中国东南部和中部的多个省份^[3−4]；山蜡梅仅在长江以南有较窄的分布^[5]；柳叶蜡梅主要生长在江西、安徽南部和浙江^[6]。蜡梅属植物具有镇咳、抗炎、解热、抗菌和降压作用，花可用于提取精油，根、茎和叶可入药^[7−9]，其干燥的叶被用于治疗感冒和流感^[10−13]。其中，柳叶蜡梅和山蜡梅茎叶中的次生代谢产物，如黄酮类、香豆素类、生物碱类和甾体类被认为是药理作用的有效成分^[14−15]。

植物种类不同，其花香物质的种类与含量也不同^[16]。蜡梅花香的主要物质为萜类化合物(单萜和倍半萜)和苯环类化合物^[17−18]，山蜡梅和柳叶蜡梅干花的主要花香物质为α-月桂烯、桉叶素和四甲基环癸二烯甲醇^[19]。蜡梅在一天的不同时段内释放的花香物质，即日释放节律存在波动。盛开的磬口蜡梅C. praecox var. grandiflorus在13:00释放的花香物质种类多于乔种蜡梅C. praecox var. intermedius^[20]。在不同昼夜节律下，磬口蜡梅释放的主要花香物质种类接近，但其数量在12:00最多^[21]。

由于蜡梅属植物不同物种以及不同时间段内的花香物质存在差异，有必要对其进行系统的比较分析，以揭示不同物种间以及不同释放节律下花香物质的异同。本研究采用顶空-固相微萃取(HS-SPME)结合气相色谱-质谱联用技术(GC-MS)对3种蜡梅属植物(蜡梅、柳叶蜡梅和山蜡梅)的花香物质进行鉴定和分析，旨在比较3种蜡梅属植物的花香物质及其白天的释放节律。

3. 讨论

3.1. 3种蜡梅属植物花香物质的种类与相对含量

本研究在蜡梅、柳叶蜡梅和山蜡梅花中共检测出44种花香物质。其中，从不同节律的蜡梅花中鉴定出19种花香物质，主要为芳樟醇和乙酸苄酯，这2种物质在其他针对蜡梅花香变化的研究中也被证明大量存在^[22−24]，这进一步验证了其在蜡梅花香中的核心作用。另外，蜡梅的主要花香成分包括烯类、醇类和酯类化合物，这与他人研究结果一致^[25]。目前有关柳叶蜡梅和山蜡梅鲜花花香物质的研究较少，只在其干花中鉴定出α-月桂烯^[26]。本研究表明柳叶蜡梅花中含量最多的花香物质是β-水芹烯，β-水芹烯和(+)-柠檬烯在山蜡梅的花中也占有很大比例。

3.2. 3种蜡梅属植物花香物质的差异

从蜡梅、柳叶蜡梅和山蜡梅中分别鉴定出19、23和18种花香物质。结果表明3种蜡梅属植物之间的花香物质差异较大。3种蜡梅属植物鉴定出的花香物质中，只有β-水芹烯和β-石竹烯是3个物种共有的，多达30种物质仅在其中一个物种中检测到。仅存在于蜡梅花中的物质(15种)比仅存在于柳叶蜡梅(9种)和山蜡梅(6种)花中的物质多。2种花香物质(芳樟醇和β-月桂烯)在蜡梅和柳叶蜡梅花中存在，而在山蜡梅花中未检测到；10种物质是柳叶蜡梅和山蜡梅花共有的，而在蜡梅花中未检测到。

在9类花香物质中，烯类、醇类和酯类是种类与含量最丰富的类别。物种间的差异性与多样性也表现在这3个类别中。从3种蜡梅属植物中检测到15种烯类化合物，蜡梅中仅检测出5种，而柳叶蜡梅和山蜡梅花中分别检测到9和10种。只有2种烯类(β-水芹烯和β-石竹烯)为3个物种所共有。共检测到13种醇类化合物，蜡梅和柳叶蜡梅花中分别检测到8和6种，而山蜡梅花中仅检测到2种。其中，芳樟醇为蜡梅与柳叶蜡梅共有，4-萜烯醇和α-松油醇为柳叶蜡梅和山蜡梅共有。共检测到8种酯类化合物，其中2种为柳叶蜡梅和山蜡梅共有，3种为蜡梅特有，3种为柳叶蜡梅特有。

柳叶蜡梅和山蜡梅的花香物质含量较为相似，但蜡梅与其他2种植物的花香物质相对含量差异较大。在柳叶蜡梅和山蜡梅中，β-水芹烯是相对含量最丰富的花香物质，而在蜡梅中并未检测到。在蜡梅中，芳樟醇(49.14%~59.16%)与乙酸苄酯(17.67%~24.34%)是相对含量最高的花香物质，而柳叶蜡梅和山蜡梅花中不含芳樟醇与乙酸苄酯。这可能与系统发育关系有关。系统发育分析表明，柳叶蜡梅和山蜡梅的亲缘关系比蜡梅更近^[27]。另一方面，作为常见的花香物质，芳樟醇和乙酸苄酯对传粉昆虫具有一定的吸引作用^[28−29]。鉴于蜡梅特殊的低温开花环境，可以推测蜡梅的传粉昆虫影响其花香物质的释放。

3.3. 3种蜡梅属植物花香物质的释放节律

3种蜡梅属植物花香物质的白天释放节律存在差异。柳叶蜡梅和山蜡梅花在上午、中午和下午的花香物质差异较大，而蜡梅的花香物质在3个时间段内变化相对较小。本研究显示：蜡梅在中午检测出的花香物质种类最多，这与先前关于蜡梅花香释放节律的报道结果一致^[24]。同样，柳叶蜡梅也在中午释放出种类最为丰富的花香物质。然而，山蜡梅在3个时间段内检测出的花香物质的种类数接近，上午检测到的花香物质种类较多。相关研究揭示了矮牵牛Petunia hybrida、烟草Nicotiana suaveolens等多种园艺植物释放花香物质的昼夜节律模式^[30−31]。矮牵牛作为模式物种被用于研究植物花香物质的释放节律^[32]，其在夜间散发出强烈的香味，这可能与传粉综合症有关。矮牵牛花香物质的释放遵循内源性昼夜节律，大约在黑暗4~6小时后达到峰值^[33−34]。由此可见，温度、湿度与光照强度可能对花香物质的释放产生了影响。一般而言, 植物挥发性物质随着温度、光照等强度的增加而增加^[35]。

4. 结论

从3种蜡梅属植物中共鉴定出44种花香物质，蜡梅、柳叶蜡梅和山蜡梅中分别鉴定出19、23和18种花香物质。这44种物质被划分为9类，其中烯类、醇类和酯类是种类与含量最丰富的类别。通过比较分析可知，柳叶蜡梅和山蜡梅花香物质较为相似，验证了两者在系统发育上的关联性。蜡梅的主要花香物质为芳樟醇和乙酸苄酯，与其他2种植物的花香物质种类差异较大。由于蜡梅特殊的低温开花环境，其传粉昆虫与花香之间的相互作用可能是花香物质产生独特性的原因。本研究揭示了蜡梅属3个物种花香物质的异同，为蜡梅属植物遗传育种、资源利用及开发更深层次的价值提供理论基础。

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物质类别	物质名称	保留指数	相对含量/%
物质类别	物质名称	保留指数	蜡梅	柳叶蜡梅	山蜡梅
烯类	莰烯 Camphene	964	−/−/−	2.31±0.22/2.32±0.08/ 2.71±0.33	1.04±0.25/−/ 1.63±0.06
	(-)-莰烯 L-Camphene	965	−/−/−	−/−/−	−/0.83±0.22/−
	β-水芹烯 β-Phellandrene	989	0.40±0.11/−/−	80.65±8.97/77.49±4.52/ 79.98±4.54	54.96±5.58/42.02±3.59/ 55.50±1.46
	β-蒎烯 β-Pinene	1 006	−/−/−	−/2.95±0.72/ 6.43±0.62	5.00±0.77/2.29±0.64/ 5.43±0.64
	月桂烯 β-Myrcene	1 007	0.99±0.15/1.01±0.05/ 1.02±0.03	4.91±1.42/−/−	−/−/−
	4-蒈烯 (+)-4-Carene	1 027	−/−/−	1.09±0.31/0.70±0.17/ 1.22±0.04	0.67±0.13/1.09±0.26/ 1.38±0.16
	(+)-柠檬烯 D-Limonene	1 039	−/−/−	−/−/−	30.67±7.26/47.41±6.08/ 31.70±7.57
	(E)-Β-罗勒烯 trans-β-Ocimene	1 048	−/−/−	−/−/0.06±0.01	−/−/−
	γ-松油烯 γ-Terpinene	1 067	−/−/−	2.49±0.45/1.88±0.08/ 2.21±0.07	1.54±0.12/2.65±0.40/ 2.46±0.29
	别罗勒烯 Allocimene B	1 135	1.26±0.23/1.36±0.06/ 1.31±0.03	−/−/−	−/−/−
	(-)-α-荜澄茄油烯 α-Cubebene	1 352	−/−/−	−/−/−	−/−/0.08±0.02
	毕澄茄烯 Cubebene	1 392	−/−/−	−/−/−	−/−/0.11±0.02
	β-榄香烯 β-Elemen	1 394	−/0.09±0.03/−	−/−/−	−/−/−
	β-石竹烯 Caryophyllene	1 425	0.17±0.02/0.30±0.07/ 0.13±0.02	0.21±0.02/0.12±0.03/ 0.11±0.04	0.10±0.03/−/ 0.31±0.03
	Δ-杜松烯 δ-Cadinene	1 513	−/−/−	−/0.09±0.03/ 0.24±0.06	−/−/−
醇类	苯甲醇 Benzyl alcohol	1 044	10.76±2.02/7.49±0.54/ 5.05±0.58	−/−/−	−/−/−
	顺-Α,Α-5-三甲基-5-乙烯基四氢化呋喃-2-甲醇 2-Furanmethanol, 5-ethenyltetrahydro-α,α,5-trimethyl-, cis-	1 096	0.60±0.24/−/ 0.53±0.08	−/−/−	−/−/−
	反式-4-(异丙基)-1-甲基环己-2-烯-1-醇 2-Cyclohexen-1-ol, 1-methyl-4-(1-methylethyl)-, trans-	1 103	−/−/−	2.48±0.76/−/−	−/−/−
	芳樟醇 Linalool	1 112	49.14±8.82/55.59±2.81/ 59.16±3.64	−/0.41±0.12/−	−/−/−
	苯乙醇 Phenylethyl Alcohol	1 117	−/−/−	−/0.55±0.21/ 0.78±0.28	−/−/−
	冰片 endo-Borneol	1 167	−/−/−	0.83±0.20/−/−	−/−/−
	桃金娘烯醇 Myrtenol	1 174	−/0.06±0.01/−	−/−/−	−/−/−
	2,2,6-三甲基-6-乙烯基四氢-2H-呋喃-3-醇 2H-Pyran-3-ol, 6-ethenyltetrahydro-2,2,6-trimethyl-	1 178	−/0.09±0.01/ 0.12±0.01	−/−/−	−/−/−
	4-萜烯醇 Terpinen-4-ol	1 179	−/−/−	−/−/0.14±0.02	0.17±0.04/−/−
	α-松油醇 α-Terpineol	1 192	−/−/−	2.06±0.20/7.31±2.76/ 1.80±0.25	4.05±1.87/0.24±0.05/ 0.69±0.28
	橙花醇 cis-Geraniol	1 260	0.07±0.02/0.11±0.01/ 0.07±0.02	−/−/−	−/−/−
	2,6-二甲基辛-2,7-二烯-1,6-二醇 2,7-Octadiene-1,6- diol, 2,6-dimethyl-	1 278	−/0.11±0.01/ 0.05±0.01	−/−/−	−/−/−
	肉桂醇 Cinnamyl alcohol	1 306	0.26±0.06/0.41±0.04/ 0.35±0.03	−/−/−	−/−/−
酯类	乙酸苄酯 Benzyl acetate	1 172	24.34±1.73/19.33±0.47/ 17.67±0.78	−/−/−	−/−/−
	水杨酸甲酯 Methyl salicylate	1 197	7.37±1.77/8.60±0.79/ 9.86±1.16	−/−/−	−/−/−
	乙酸异龙脑酯 Isobornyl acetate	1 288	−/−/−	−/0.11±0.03/−	−/−/−
	癸酸甲酯 Decanoic acid, methyl ester	1 328	−/−/−	−/0.18±0.05/−	−/−/−
	二乙二醇丁醚醋酸酯 Ethanol, 2-(2-butoxyethoxy)-, acetate	1 371	−/−/−	0.28±0.04/ 0.40±0.12/−	0.32±0.13/ 0.06±0.02/−
	2-癸烯酸甲酯 2-Decenoic acid, methyl ester	1 374	−/−/−	−/0.33±0.08/−	−/−/−
	2,4-葵二烯酸甲酯 2,4-Decadienoic acid, methyl ester, (E,Z)-	1 397	−/−/−	0.97±0.33/2.81±0.16/ 2.62±0.44	0.29±0.05/−/−
	乙酸桂酯 Cinnamyl acetate	1 447	−/0.04±0.01/−	−/−/−	−/−/−
酚类	丁香酚 Eugenol	1 360	1.20±0.21/1.26±0.12/ 0.99±0.19	−/−/−	−/−/−
醛类	苯甲醛 Benzaldehyde	977	0.15±0.02/0.12±0.00/ 0.11±0.00	−/−/−	−/−/−
	2-(4-甲基-3-环己烯-1-基)丙醛 3-Cyclohexene-1- acetaldehyde, α,4-dimethyl-	1 176	−/−/−	−/−/−	0.96±0.19/2.55±0.78/ 0.40±0.14
杂环类	吲哚 Indole	1 294	3.30±0.23/4.03±0.09/ 3.60±0.20	−/−/−	−/−/−
芳香烃类	对二甲苯 P-Xylene	915	−/−/−	1.00±0.27/1.82±0.54/ 1.12±0.32	−/0.46±0.12/ 0.18±0.05
酮类	樟脑 Camphor	1 147	−/−/−	−/0.09±0.02/ 0.12±0.04	−/−/−
其他	(+)-氧化柠檬烯 (+)-Limonene oxide	1 137	−/−/−	0.74±0.13/0.45±0.02/ 0.46±0.04	0.19±0.02/0.41±0.12/ 0.12±0.04
	石竹素 Caryophyllene oxide	1 533	−/−/−	−/−/−	0.05±0.01/−/−
总计	44		14/17/15	13/18/15	14/11/13
说明：相对含量的3个数据依次在9:00/12:00/15:00检测，数值为平均值±标准差。−表示未检出。

物质类别	蜡梅		柳叶蜡梅		山蜡梅		物质类别	蜡梅		柳叶蜡梅		山蜡梅
物质类别	种数	小计	种数	小计	种数	小计	物质类别	种数	小计	种数	小计	种数	小计
烯类	4/4/3	5	6/7/8	9	7/6/9	10	杂环类	1/1/1	1	−/−/−	−	−/−/−	−
醇类	5/7/7	8	3/3/3	6	2/1/1	2	芳香烃类	−/−/−	−	1/1/1	1	−/1/1	1
酯类	2/3/2	3	2/5/1	5	2/1/−	2	酮类	−/−/−	−	−/1/1	1	−/−/−	−
酚类	1/1/1	1	−/−/−	−	−/−/−	−	其他	−/−/−	−	1/1/1	1	2/1/1	2
醛类	1/1/1	1	−/−/−	−	1/1/1	1	合计	14/17/15	19	13/18/15	23	14/11/13	18
说明：种数的3个数据依次在9:00/12:00/15:00检测。−表示未检出。

类别	相对含量/%
类别	蜡梅	柳叶蜡梅	山蜡梅
烯类	2.82/2.76/2.46	91.66/85.55/92.96	93.98/96.29/98.60
醇类	60.83/63.86/65.33	5.37/8.27/2.72	4.22/0.24/0.69
酯类	31.71/27.97/27.53	1.25/3.83/2.62	0.61/0.06/−
酚类	1.20/1.26/0.99	−/−/−	−/−/−
醛类	0.15/0.12/0.11	−/−/−	0.96/2.55/0.40
杂环类	3.30/4.03/3.60	−/−/−	−/−/−
芳香烃类	−/−/−	1.00/1.82/1.12	−/0.46/0.18
酮类	−/−/−	−/0.09/0.12	−/−/−
其他	−/−/−	0.74/0.45/0.46	0.24/0.41/0.12
说明：相对含量的3个数据依次在9:00/12:00/15:00检测。−表示未检出。