Effect of drought stress and re-watering on emissions of volatile organic compounds from <i>Rosmarinus officinalis</i>

LIU Fang; XU Gaiping; WU Xingbo; DING Qianqian; ZHENG Jie; ZHANG Rumin; GAO Yan

doi:10.11833/j.issn.2095-0756.2014.02.015

Volume 31 Issue 2

Mar. 2014

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Article Navigation > Journal of Zhejiang A&F University > 2014 > 31(2): 264-271

LIU Fang, XU Gaiping, WU Xingbo, DING Qianqian, ZHENG Jie, ZHANG Rumin, GAO Yan. Effect of drought stress and re-watering on emissions of volatile organic compounds from Rosmarinus officinalis[J]. Journal of Zhejiang A&F University, 2014, 31(2): 264-271. doi: 10.11833/j.issn.2095-0756.2014.02.015

Citation:

LIU Fang, XU Gaiping, WU Xingbo, DING Qianqian, ZHENG Jie, ZHANG Rumin, GAO Yan. Effect of drought stress and re-watering on emissions of volatile organic compounds from Rosmarinus officinalis[J]. Journal of Zhejiang A&F University, 2014, 31(2): 264-271. doi: 10.11833/j.issn.2095-0756.2014.02.015

Effect of drought stress and re-watering on emissions of volatile organic compounds from Rosmarinus officinalis

doi: 10.11833/j.issn.2095-0756.2014.02.015

School of Forestry and Biotechnology, Zhejiang A & F University, Lin'an 311300, Zhejiang, China

Received Date: 2013-01-16
Rev Recd Date: 2013-05-18
Publish Date: 2014-04-20

Abstract

To better understand the effects of water limitations on the emission of volatile organic compounds (VOCs), VOC emissions from Rosmarinus officinalis were determined under drought stress and after re-watering. Results showed that with drought stress R. officinalis VOC emissions and lipoxidase activity were strongly affected with the degree of influence dependent on the stress level and time. At the beginning of the drought stress period, lipoxidase activity increased slightly; midway through, the activity reached a peak; then it declined. Drought stress reduced R. officinalis VOC emissions, but induced new VOCs including 2-hexenal, leaf alcohol, sorbaldehyde, and n-decanal. Also, R. officinalis increased terpene emissions, especially monoterpenes, to reduce drought stress damage.
- botany,
- drought stress,
- re-water,
- Rosmarinus officinalis,
- lipoxidase,
- volatile organic compounds (VOCs),
- green leaf volatiles (GLVs),
- monoterpenes

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园林植物是维系城市生态平衡的重要因素,自然环境的干旱胁迫会使植物体内细胞脱水,造成水分亏缺而导致植物受害^[1]。干旱胁迫是众多环境胁迫因子中最常见的一种^[2],对植物的生长发育会产生不同程度的影响。在现代化的城市绿化中,选择观赏性好、抗性强的观赏植物,对节约水资源改善土壤干旱、半干旱的现状具有重要意义。植物挥发性有机化合物(volatile organic compounds,简称VOCs)通常指分子质量小于250,沸点小于340 ℃,从植物地上部分(如叶、花和芽等)表面散发的多种微浓度的挥发性次生代谢物质^[3]。植物VOCs在生态系统中占有重要地位,它是重要的化学信息传递物质^[4-5],在调节植物的生长、发育和繁衍,抵御环境胁迫及预防动物和昆虫的危害等^[3-10]方面具有重要作用。非生物胁迫会明显影响植物VOCs的合成与释放,Ibrahim等^[11]和Hartikainen等^[12]均研究发现,高温胁迫会诱导欧洲白桦Betula pendula和欧洲山杨Populus tremula释放大量的异戊二烯、萜类和C6绿叶性气体(green leaf volatiles,GLVs)；光照—黑暗转换处理会诱导欧洲黑杨Populus deltoides叶片大量释放己烯醛和乙醛等^[13]。干旱对VOCs释放规律的影响呈现多样性,Staudt等^[14]在对栓皮栎Quercus suber的研究中发现,适度干旱胁迫导致单萜类化合物的释放量增加30%,而Lavoir等^[15]发现,干旱胁迫下冬青栎Quercus ilex。释放的单萜是对照处理的55%。Blanch等^[16]研究发现干旱导致冬青栎萜烯类释放量增加33%,而相同干旱条件下地中海白松Pinus halepensis萜烯类释放量降低了38%。迷迭香 Rosemarinus offcinalis,又名情人草,原产地为欧洲、北非及地中海沿岸,主要分布在法国、突尼斯、摩洛哥和保加利亚。迷迭香枝叶气味浓烈,抗旱性极强,是一种被广泛应用的园林芳香植物,可用于花坛绿地片植、丛植、孤植或作为配材,用于镶边,亦可用作小绿篱或花篱,国外多用于花境配材。近几年,迷迭香的研究多集中在其有效成分提取工艺^[17]、抗氧化物功能^[18]和精油组分分析^[19]等方面。关于干旱胁迫下迷迭香VOCs释放规律与抗旱机制之间的关系未见报道。因此,本研究采用称量控水法模拟干旱胁迫,利用动态顶空气体循环法和热脱附-气相色谱-质谱联用技术(TDC-GC-MS)测定了干旱胁迫及复水后迷迭香VOCs成分的变化；同时,测定了迷迭香叶片脂氧合酶(LOX)活性的变化,以期从迷迭香释放VOCs角度揭示它对干旱胁迫的适应机制,为进一步研究植物释放VOCs对非生物胁迫的响应机制提供理论依据。

1. 材料与方法

1.1. 供试材料及处理

供试材料为迷迭香2年生实生苗(江苏省宿迁谊园花卉盆景苗木公司提供),苗高约为20 cm。2012年4月栽置于盛有培养土[m(泥炭): m(沙土): m(蛭石)=1: 2: 1,干质量1.8 kg]的花盆中(盆径为18 cm,盆高为20 cm),1株·盆^-1,盆栽苗置于温室中,在充足的水分条件下培养恢复1个月后实施控水处理。选取长势一致、枝条数相同的盆栽苗24盆,分别进行干旱胁迫处理,设正常浇水(对照,75%田间持水量)、轻度干旱(60% 田间持水量)、中度干旱(40% 田间持水量)和重度干旱(20% 田间持水量)等4种处理,6盆·处理^-1。采用称量法控制土壤水分含量,每天傍晚进行称量并补充各处理消耗的水分。4 d测定1次脂氧合酶活性,重复3次。干旱胁迫处理24 d后,对重度干旱进行复水处理,每天测定脂氧合酶活性,复水共6 d。

1.2. 实验方法

酶液提取：称取迷迭香枝条第5~8片叶片0.5 g,加入适量预冷的0.05 mol·L^-1的pH 7.8磷酸缓冲溶液,充分冰浴研磨后,定容至25.0 mL。4 ℃下1万r·min^-1离心20 min,回收上清液用于脂氧合酶活性测定,重复3次。

脂氧合酶活性的测定参照Hatanaka等^[20]的方法。底物配制：将0.1 mL吐温20加入到2.0 mL 0.2 mol·L^-1的硼酸缓冲液(pH 9.0)中并混匀,滴加0.1 mL亚油酸后充分混匀,再加入0.2 mL 1.0 mol·L^-1 氢氧化钠溶液并保证此时的溶液澄清,然后加入上述硼酸缓冲液18.0 mL,再加水定容至50.0 mL,-20 ℃保存备用。3.0 mL反应体系中含有0.1 mL底物、2.8 mL磷酸缓冲液(0.1 mol·L^-1,pH 7.0)和0.1 mL酶液,测定反应体系3 min内在234 nm波长处吸光度的变化。酶活性定义：234 nm波长处,以亚油酸为底物,3.0 mL反应体系吸光度1 min增加0.1为1个酶活单位。

1.3. VOCs采集及分析方法

VOCs的采集：干旱胁迫处理24 d及复水第6天,选取株型相似、枝条数相同的迷迭香,3盆·处理^-1,采用QC-2型大气采样仪通过动态顶空气体循环采集法收集VOCs,气体流量0.1 m³·min^-1,采样时间20 min。采用TDS-GC-MS分析迷迭香VOCs成分。仪器及参数设置条件参照Gao等^[21]的方法。

TDS(TD3型,GERSTEL,德国)工作条件：载气压力20 kPa；进样口温度250 ℃；热脱附温度250 ℃,10 min；冷阱温度-100 ℃,3 min；进样时冷阱骤然升至260 ℃。

GC(7890A型,Agilent,美国)工作条件：色谱柱为30 m × 250 μm × 0.25 μm的HP-5MS柱；升温程序：初始温度40 ℃,保持4 min后,以6 ℃·min^-1速率升至250 ℃,保持3 min后,以10 ℃·min^-1的速率升至270 ℃,保持5 min。

MS(5975C型,Agilent,美国)工作条件：电力方式为EI,电子能量为70 eV,质量范围为28~450,接口温度为280 ℃,离子源温度为230 ℃,四级杆温度为150 ℃。

1.4. 数据分析

采用NIST2008谱库检索,采用峰面积表示VOCs成分的相对含量。使用Oringin 8.0和SPSS软件进行数据处理、图表制作及统计分析,方差分析采用Duncan新复极差法。

3. 讨论和结论

脂氧合酶通过催化具有顺,顺-1,4-戊二烯结构的不饱和脂肪酸的加氧反应,生成含氧挥发性有机化合物(oxygenated volatile hydrocarbons,OVOCs),其中包括绿叶性气体(GLVs)^[3]。动物取食、病原菌侵害和机械损伤均能诱导植物大量释放GLVs^[22-23],这些化合物都具有提高植物受损伤器官抵御细菌侵害的功能,可以作为植物体内的信号物质来增加防御,提高植物体的抗逆性^[24-25]。干旱胁迫诱导产生大量活性氧(reactive oxygen species,ROS),活性氧破坏细胞膜结构,使得细胞内游离不饱和脂肪酸增加,进而促使脂氧合酶活性升高,催化产生大量的OVOCs。本研究发现,随着干旱胁迫的进行,各处理的脂氧合酶活性均增加。在对挥发性有机化合物(VOCs)各组分进行分析时发现,干旱诱导产生了3种醛类物质和2种C6-GLVs,这些物质的相对含量增加,这与脂氧合酶活性增加呈现相关性；并且轻度干旱和中度干旱处理的3-己烯-2-酮含量明显高于对照,此时这2个处理的脂氧合酶活性也高于对照,两者存在相关性；复水后,脂氧合酶活性降低,干旱胁迫诱导产生的2-己烯醛、山梨醛和癸醛也同时消失。轻度干旱处理酯类相对含量是对照的1.66倍,而中度干旱和重度干旱处理的酯类相对含量均下降,可能是因为轻度干旱下脂氧合酶活性最高,催化生成大量酯类,另一种可能的原因是,迷迭香体内具有储存VOCs的结构^[26],轻度干旱下酯类物质储存结构中释放出来,而中度干旱和重度干旱处理储存的VOCs在干旱早期或已经释放,具体原因有待于进一步研究。

萜烯类化合物是植物次生代谢产生的种类最多、释放量最大的VOCs,对植物体起到重要的保护作用^{[22, 27]}。本研究发现各干旱处理的迷迭香萜烯类百分含量均有较大增加,一方面是因为萜类化合物沸点较低,干旱诱导气孔关闭时,它们可以通过非气孔途径向外界排放^[28]；另一方面,当非生物胁迫发生时,植物体内会通过增加萜烯类化合物的释放来缓解胁迫压力^{[3, 27-28]}。Simpraga等^[29]在对欧洲山毛榉Fagus sylvatica的研究中发现,停止浇水10 d后,山毛榉净光合速率下降,几天之后单萜类的释放总量减少,但分配到单萜合成途径中碳的比例却是增加的,说明植物体能够通过增加单萜类的释放来降低光合氧化伤害^[30]；Lavoir等^[15]发现：干旱胁迫下冬青栎释放的单萜是对照处理的55%。本研究同样表明：各干旱处理单萜类释放总量均减少,但单萜类相对含量却随着干旱程度的加深呈现明显的增加趋势,说明迷迭香能够通过增加单萜类的释放来抵御干旱胁迫。

综上所述：干旱能够诱导脂氧合酶酶活性增加,催化合成更多的GLVs及醛类VOCs；干旱导致迷迭香VOCs释放总量下降,却诱导迷迭香合成新的VOCs成分,且萜烯类的释放比例呈增加的趋势。研究表明：迷迭香能够通过增加GLVs、醛类和萜烯类化合物的释放来有效缓解干旱胁迫造成的伤害,提高迷迭香的抗旱性。园林植物在生长发育的过程中,不可避免的会受到环境的胁迫,而干旱胁迫是最常见的一种,研究园林植物的抗旱性机理对于园林植物的养护管理意义重大,并且,迷迭香VOCs的释放规律研究可以为迷迭香的园林配置,尤其是芳香植物园配置提供科学依据。

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t/min	挥发性有机物	分子式	分子量	峰面积A×10⁶
t/min	挥发性有机物	分子式	分子量	对照	轻度干旱	中度干旱	重度干旱	复水
5.18	3-己烯-2-酮 3-hene-2-one	C₆H₁₀O	98	2.47 ± 0.03	8.58 ± 0.01	5.40 ± 0.11	1.40 ± 0.01	2.28 ± 0.01
6.66	2-己烯醛（E)-2-hexenal	C₆H₁₀O	98	-	-	1.03 ± 0.01	1.09 ± 0.02	-
6.78	叶醇 leaf alcohol	C₆H₁₂O	100	-	1.07 ± 0.02	1.17 ± 0.02	0.89 ± 0.01	0.63 ± 0.01
7.42	α-薇烯α-pinene	C₁₀H₁₆	136	3.79 ± 0.03	0.78 ± 0.02	3.83 ± 0.10	1.66 ± 0.11	1.40 ± 0.04
8.19	山梨醛 sorba ehyde	C₆H₈O	96	-	-	-	0.51 ± 0.00	-
8.47	三环烯 tricyclene	C₁₀H₁₆	136	-	-	0.56 ± 0.00	0.74 ± 0.01	0.70 ± 0.01
8.69	侧柏烯thujene	C₁₀H₁₆	136	9.68 ± 0.23	7.74 ± 0.41	6.64 ± 0.21	6.02 ± 0.12	7.71 ± 0.19
8.91	β薇烯β-pinene	C10H16	136	79.85 ± 3.44	48.72 ± 2.68	73.56 ± 4.95	76.26 ± 6.18	78.93 ± 5.36
9.24	樟烯 camphene	C₁₀H₁₆	136	16.82±1.11	8.82± 0.89	16.41±1.55	15.13±2.02	16.83±2.58
9.39	伞柳醇 umbellulol	C₁₀H₁₆O	l52	l2.50±l.87	15.12±2.07	9.64±1.01	6.56±0.66	7.99±0.89
10.00	桧稀 sabinene	C₁₀H₁₆	136	15.46±1.29	9.52±1.06	14.30±2.03	12.48±1.69	14.21±1.87
10.52	月桂烯myrcene	C₁₀H₁₆	136	50.11±3.48	53.59±4.08	26.28±2.22	14.81±1.23	19.90±2.10
10.80	水芹稀 phellandrene	C₁₀H₁₆	l36	l8.93±l.07	l7.50±2.04	8.78±0.67	4.64±0.2l	6.79±0.24
10.88	蒈烯 3-carene	C₁₀H₁₆	l36	-	-	1.19±0.02	0.79±0.00	0.75±0.00
11.12	α-萜品烯 α-terpinene	C₁₀H₁₆	136	23.81±2.34	21.41±2.45	10.65±1.67	5.49±0.19	8.16±0.98
11.37	伞花经o-cymene	C₁₀H₁₄	134	-	93.77±7.33	55.02±3.89	19.40±1.22	36.31±1.32
11.56	a-萜品醇 a-terpineol	C₁₀H₁O	154	96.79±7.92	7.80±0.11	1.08±0.01	20.77±2.03	25.12±1.98
11.99	罗勒烯ocimene	C₁₀H₁₆	136	4.82±0.66	11.63±0.98	6.35±0.93	3.51±0.06	5.48±0.08
12.35	β萜品烯β-terpinene	C₁₀H₁₆	136	50.44±4.22	52.84±3.98	19.77±1.76	10.44±1.05	16.44±1.08
13.08	萜品油烯terpinolene	C₁₀H₁₂	132	32.63±2.87	33.23±3.48	15.09±0.94	7.91±0.56	11.70±1.86
13.44	里那醇linalool	C₁₀H₁₈O	154	32.06±3.04	29.88±2.65	15.82±1.44	6.08±0.72	9.43±0.57
13.59	蒿萜酮eucarvone	C₁₀H₁₆O	140	3.42±0.22	2.25±0.43	1.71±0.18	0.43±0.01	8.64±1.00
14.07	菊油环丽 chrysanthenone	C₁₀H₁₄O	150	39.76±2.94	29.38±1.69	17.32±1.03	6.31±0.08	7.23±0.11
14.39	松香芹醇pinocarveol	C₁₀H₁₆O	152	1.11±0.02	1.78±0.03	1.46±0.01	0.62±0.00	1.02±0.02
14.53	棒脑 camphor	C₁₀H₁₆O	152	26.99±1.93	19.95±2.03	20.43±1.49	7.91±0.99	11.96±1.02
14.74	异龙脑 isoborneol	C₁₀H₁₈O	154	0.56±0.00	0.56±0.01	0.71±0.01	20.20±2.03	0.54±0.00
14.93	松香芹酮pinocarvone	C₁₀H₁₄O	150	2.12±0.02	4.71±0.08	3.36±0.05	1.26±0.11	2.12±0.12
15.21	龙脑 borneol	C₁₀H₁₈O	154	44.98±2.87	39.91±3.18	47.86±3.96	2.81±0.02	29.22±2.77
15.28	松莰酮 pinocamphone	C₁₀H₁₆O	152	4.18±0.44	1.13±0.04	3.18±0.62	2.94±0.17	3.78±0.32
15.34	4-萜品醇 4-carvomenthenol	C₁₀H₁₈O	154	10.84±0.87	12.45±1.03	7.97±0.06	8.34±0.33	4.85±0.11
15.64	2-薇烯-4-酮 2-pinen-4-one	C₁₀H₁₄O	150	1.09±0.09	-	0.91±0.00	2.86±0.11	-
15.81	草蒿脑estragole	C₁₀H₁₂O	148	-	-	2.04±0.04	3.22±0.05	3.30±0.03
15.87	顺式萜品醇cis-terpineol	C₁₀H₁₈O	154	17.34±1.22	14.03±1.83	9.11±0.79	-	6.98±0.77
16.03	癸醒 n-decanal	C₁₀H₂₀O	156	-	-	1.32±0.22	-	-
16.06	桃金娘烯醇myrtenol	C₁₀H₁₆O	152	3.14±0.19	4.94±0.36	1.62±0.09	-	-
16.28	反式马鞭草烯酮 rans - verbenone	C₁₀H₁₄O	150	24.59±1.43	24.21±2.64	34.97±2.92	21.18±1.88	26.42±1.59
16.33	2-烯丙基-4-甲基苯酚 2-allyl-4-methylphenol	C₁₀H₁₂O	148	-	-	0.91±0.00	3.22±0.11	0.90±0.01
16.77	香茅醇 citronellol	C₁₀H₂₀O	198	2.13±0.23	2.78±0.09	1.21±0.11	0.79±0.01	1.16±0.04
16.92	百里香酚甲醚 ethylthymylether	C₁₀H₁₆O	164	-	0.48±0.00	-	-	-
17.08	蒈醇 caranol	C₁₀H₁₈O	154	1.75±0.12	1.28±0.07	0.96±0.06	-	0.44±0.00
17.29	桃金娘院醇myrtanol	C₁₀H₁₆O	136	4.17±0.11	4.01±0.21	2.76±0.22	-	1.44±0.02
17.70	香叶醇lemonol	C₁₀H₁₈O	154	8.37±0.17	12.45±0.88	4.44±0.65	1.98±0.06	2.24±0.06
18.27	梓檬醒guaniol	C₁₀H₁₆O	152	3.52±0.33	17.39±0.32	0.84±0.00	2.29±0.17	1.77±0.03
18.34	香芹酮carvone	C₁₀H₁₄O	150	3.31±0.21	-	2.94±0.07	2.34±0.08	1.80±0.07
18.41	菊酸甲酯 methyl chrysanthemate	C₁₀H₁₈O₂	182	1.43±0.04	3.94±0.15	1.31±0.06	0.56±0.00	0.92±0.02
18.87	乙酸龙脑酯bornyl acetate	C₁₀H₂₀O₂	196	44.36±3.77	70.24±7.86	22.72±2.83	14.18±1.93	21.38±1.77
19.06	菊酸乙酯 chrysanthenyl acetate	C₁₂H₁₈O₂	194	0.74±0.01	1.31±0.10	1.03±0.07	-	-
20.06	乙酸桃金娘烯酯 myrtenyl acetate	C₁₂H₁₈O₂	194	1.46±0.10	2.76±0.21	0.65±0.02	-	0.49±0.01
20.35	冰片烯bornylene	C₁₀H₁₆	136	1.28 ± 0.04	2.96 ± 0.05	1.27 ± 0.01	0.49 ± 0.00	0.70 ± 0.04
20.90	香茅醇乙酸酯 citronellyl acetate	C₁₀H₁₈	138	0.49 ± 0.03	6.75 ± 0.31	0.43 ± 0.02	-	0.53 ± 0.01
21.23	乙酸异洋薄荷酯 isopulegol acetate	C₁₀H₁₈O	154	3.32 ± 0.21	1.61 ± 0.01	0.61 ± 0.01	0.41 ± 0.00	0.58 ± 0.01
21.82	乙酸香叶酯 geraniol acetate	C₁₀H₁₄O	150	2.64 ± 0.20	2.78 ± 0.25	0.90 ± 0.02	-	1.00 ± 0.11
22.20	顺式马鞭草烯酮 cis-verbenone	C₁₀H₁₄O	150	10.45 ± 1.03	7.40 ± 0.31	5.16 ± 0.43	0.72 ± 0.05	1.36 ± 0.07
22.46	丁香油酚甲醚 eugenol methylether	C₁₀H₁₄O₂	178	9.52 ± 0.67	13.70 ± 1.02	5.19 ± 0.48	3.03 ± 0.28	4.45 ± 0.22
22.93	反式石竹烯 trans-caryophyllene	C₁₅H₂₄	204	44.75 ± 3.89	39.11 ± 2.77	20.27 ± 1.88	8.09 ± 0.88	12.75 ± 1.11
23.64	香叶基丙酮 dihydropseudoionon	C₁₃H₂₂0	194	1.35 ± 0.21	1.07 ± 0.07	0.92 ± 0.01	-	-
23.72	顺式石竹烯 cis-caryophyllene	C₁₅H₂₄	204	6.71 ± 0.44	6.06 ± 0.28	2.83 ± 0.10	1.16 ± 0.05	1.75 ± 0.06
说明:"-"表示未检测到化合物

Effect of drought stress and re-watering on emissions of volatile organic compounds from Rosmarinus officinalis

doi: 10.11833/j.issn.2095-0756.2014.02.015