‘波叶金桂’对干旱和高温胁迫的生理生态响应

doi:10.17521/cjpe.2018.0017

摘要/Abstract

摘要：

为了探讨植物对干旱、高温及协同胁迫的响应, 该研究以木犀(Osmanthus fragrans) ‘波叶金桂’为材料, 采用盆栽质量控水法模拟干旱胁迫(对照、轻度、中度和重度)和高温胁迫, 利用动态顶空气体循环吸附法和热脱附-气相色谱-质谱(TDS-GC-MS)联用技术对其挥发性有机化合物(VOCs)进行测定; 同时测定其非结构性碳(NSC)含量及次生代谢酶活性。结果表明: 干旱胁迫对‘波叶金桂’叶片NSC组分含量影响不显著; 可溶性糖和淀粉含量在高温胁迫下显著降低, 在协同胁迫后持续性下降, 重度协同胁迫下, 葡萄糖、果糖、蔗糖和淀粉分别比对照降低47.7%、46.4%、34.4%和38.2%。干旱胁迫和协同胁迫下3-羟基-3-甲基戊二酸单酰辅酶A还原酶(HMGR)、1-脱氧木酮糖-5-磷酸还原酶(DXR)活性表现出先上升后下降, 而脂氧合酶(LOX)活性表现出持续性上升趋势; 高温胁迫后, HMGR、DXR和LOX活性显著高于对照。干旱胁迫下萜烯类VOCs释放量表现出先上升后下降趋势, 中度干旱胁迫和高温胁迫下分别比对照高37.9%和32.3%; 协同胁迫下萜烯类释放量逐渐降低, 干旱、高温和协同胁迫诱导醛类释放量明显增加。上述结果表明: 干旱胁迫条件下, ‘波叶金桂’通过NSC进行自我渗透调节, 同时合成大量萜烯类化合物来提高抗旱性; ‘波叶金桂’调控萜烯类化合物合成以及绿叶挥发物(GLVs)的释放抵御高温胁迫; 协同胁迫下萜烯类化合物的合成途径受阻, ‘波叶金桂’提高GLVs合成与释放量抵御协同胁迫; 中度和重度协同胁迫导致‘波叶金桂’细胞膜严重受损, 自我调节能力降低。

关键词: 木犀, 干旱高温胁迫, 非结构性碳, 挥发性有机化合物, 次生代谢酶

Abstract:

Aims Drought and heat stresses are two critical environmental factors affecting the growth and development of plants; climate change has exacerbated the occurrence of these conditions in many parts of the world. To elucidate the mechanisms of responses to drought and heat stresses in Osmanthus fragrans, we studied changes in nonstructural carbohydrate (NSC) and volatile organic compounds (VOCs) emissions using 3-year-old seedlings (cultivar ‘Boyejingui’) under conditions of low drought (LD), moderate drought (MD) and severe drought (SD), interactively with heat treatments (40 °C in daytime (12 h) and 30 °C at night (12 h)).

Methods The VOCs emissions were measured using the dynamic headspace air-circulation method, and the composition and content of VOCs were analyzed using the thermal desorption system/gas chromatography/mass spectrometer technique (TDS-GC-MS). We determined the content of NSC and activities of lipoxygenase (LOX), 3-hydroxy-3-methylglutaryl-CoA reductase (HMGR) and 1-deoxy-D-xylulose 5-phosphate reductoisomerase (DXR) in seedlings under different drought and heat treatments.

Important findings The content of NSC in leaves was not significantly affected by drought stress. The contents of soluble sugar and starch were significantly reduced by heat stress as well as combination of drought and heat stresses. The combination of severe drought and heat treatment significantly reduced the contents of glucose, fructose, sucrose and starch by 47.7%, 46.4%, 34.4% and 38.2% (p < 0.05), respectively. There were differences among the activities of HMGR, DXR and LOX in response to drought and heat stresses. Under drought and combined stresses, the activities of HMGR and DXR increased first and then decreased, and the LOX activity increased. The activities of HMGR, DXR and LOX were significantly higher under heat stress than in controls. The release of terpenes increased first and then decreased with the level of drought treatments. MD and SD stress stimulated the release of terpenes by 37.9% and 32.3% (p < 0.05), respectively, but the combined stresses induced a reduction in the release of terpenes. The release of aldehydes increased significantly in response to drought, heat treatment, and their combination. The results indicated that ‘Boyejingui’ seedlings can enhance resistance to drought stress by adjusting contents of non-structural carbohydrate and releasing terpene compounds. They also adjusted the biosynthesis of terpene compounds and GLVs emission to resist against heat stress. However, the biosynthesis pathway of terpene compounds was blocked in the occurrence of both drought and heat stresses, when the enhanced release of GLVs facilitated the resistance. Under heat stress, moderate and severe drought caused severe damage to cell membranes, thus reducing the self-regulatory capacity.

Key words: Osmanthus fragrans, drought and heat stress, non-structural carbohydrate, volatile organic compounds, secondary metabolic enzymes

汪俊宇, 王小东, 马元丹, 傅卢成, 周欢欢, 王彬, 张汝民, 高岩. ‘波叶金桂’对干旱和高温胁迫的生理生态响应. 植物生态学报, 2018, 42(6): 681-691. DOI: 10.17521/cjpe.2018.0017

WANG Jun-Yu, WANG Xiao-Dong, MA Yuan-Dan, FU Lu-Cheng, ZHOU Huan-Huan, WANG Bin, ZHANG Ru-Min, GAO Yan. Physiological and ecological responses to drought and heat stresses in Osmanthus fragrans ‘Boyejingui’. Chinese Journal of Plant Ecology, 2018, 42(6): 681-691. DOI: 10.17521/cjpe.2018.0017

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0017

https://www.plant-ecology.com/CN/Y2018/V42/I6/681

图/表 8

图1 干旱高温处理对波叶‘波叶金桂’叶片葡萄糖含量的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱处理的差异显著, 不同小写字母表示高温处理的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 1 Effects of drought and heat stress on glucose content in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of heat stress treatment (p < 0.05, according to LSD test). p < 0.01. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

图2 干旱高温处理对‘波叶金桂’叶片果糖含量的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 2 Effects of drought and heat stress on fructose content in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of heat stress treatment (p < 0.05, according to LSD test). **, p < 0.01. CK, control; LD, light processing; MD, moderate drought; SD, severe drought.

图3 干旱高温处理对‘波叶金桂’叶片蔗糖含量的影响(平均值±标准误差, n=5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 3 Effects of drought and heat stress on sucrose content in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of heat stress treatment (p < 0.05, according to LSD test). p < 0.01. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

图4 干旱高温处理对‘波叶金桂’叶片淀粉含量的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 4 Effects of drought and heat stress on starch content in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of heat stress treatment (p < 0.05, according to LSD test). **, p < 0.01. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

图5 干旱高温处理对‘波叶金桂’叶片3-羟基-3-甲基戊二酸单酰辅酶A还原酶活性的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 5 Effects of drought and heat stress on 3-hydroxy-3- methylglutaryl CoA reductase (HMGR) activity in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of heat stress treatment (p < 0.05, according to LSD test). **, p < 0.01. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

图6 干旱高温处理对‘波叶金桂’ 叶片1-脱氧木酮糖-5-磷酸还原酶活性的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。**, p < 0.01。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 6 Effects of drought and heat stress on 1-deoxy- D-xylulose 5-phosphate reductoisomerase (DXR) activity in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of drought and different lowercase letters denote statistically significant differences of d heat stress treatment (p < 0.05, according to LSD test). **, p < 0.01;. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

图7 干旱高温处理对‘波叶金桂’叶片脂氧合酶活性的影响(平均值±标准误差, n = 5)。不同大写字母表示干旱的差异显著, 不同小写字母表示高温的差异显著(p < 0.05, 根据LSD测验)。*, p < 0.05; **, p < 0.01;。CK, 对照; LD, 轻度干旱; MD, 中度干旱; SD, 重度干旱。

Fig. 7 Effects of drought and heat stress on lipoxygenase (LOX) activity in leaves of Osmanthus fragrans cv. ‘Boyejingui’ (mean ± SE, n = 5). Different uppercase letters indicate significant differences of different lowercase letters denote statistically significant differences of from drought and heat stress treatment (p < 0.05, according to LSD test). **, p < 0.01. CK, control; LD, light drought; MD, moderate drought; SD, severe drought.

表1 波叶金桂’叶片挥发性有机化合物的主要成分(峰面积, A × 105·10 g-1)(平均值±标准误差)

Table 1 The main components of the volatile organic compounds (VOCs) from the leaf of Osmanthus fragrans ‘Boyejingui’ (peak area, A × 105·10 g-1) (mean ± SE)

序号 No.	挥发性有机物 Volatile organic compounds	分子式 Chemical formula	28 ℃				40 ℃
序号 No.	挥发性有机物 Volatile organic compounds	分子式 Chemical formula	对照 Control	轻度干旱 Light drought	中度干旱 Moderate drought	重度干旱 Severe drought	对照 Control	轻度干旱 Light drought	中度干旱 Moderate drought	重度干旱 Severe drought
1	己烯醛 3-Hexenal	C₆H₁₀O	3.33 ± 0.23	0.58 ± 0.65	2.27 ± 0.12	-	3.61 ± 0.26	-	-	1.15 ± 0.05
2	2-甲基-4-戊烯醛 2-Methyl-4-pentenal	C₆H₁₀O	0.99 ± 0.11	2.36 ± 0.23	-	5.97 ± 0.35	-	2.88 ± 0.12	3.14 ± 0.16	1.69 ± 0.13
3	2-已烯醛 (E)-2-Hexenal	C₆H₁₀O	2.48 ± 0.18	2.14 ± 0.23	2.54 ± 0.21	8.67 ± 1.14	4.53 ± 0.54	4.35 ± 0.08	8.01 ± 0.24	7.03 ± 0.19
4	苯基丁醛 4-Phenylbutanal	C₁₀H₁₂O	-	1.85 ± 0.11	-	-	-	5.48 ± 0.13	2.58 ± 0.06	2.74 ± 0.06
5	山梨醛 Sorbaldehyde	C₆H₈O	1.08 ± 0.06	2.82 ± 0.08	1.90 ± 0.04	3.05 ± 0.64	1.80 ± 0.08	-	2.61 ± 0.11	2.33 ± 0.08
6	辛烯醛 2-Ethylhexenal	C₈H₁₄O	-	1.91 ± 0.04	4.86 ± 0.81	-	3.77 ± 0.12	1.37 ± 0.06	0.67 ± 0.02	-
7	壬醛 n-Nonanal	C₉H₁₈O	-	-	-	-	0.76 ± 0.02	1.21 ± 0.04	0.90 ± 0.05	0.53 ± 0.02
8	反-3-己烯醇 trans-3-Hexenol	C₆H₁₂O	6.80 ± 0.45	8.07 ± 0.62	7.85 ± 0.67	4.12 ± 0.35	9.26 ± 1.61	2.50 ± 0.27	5.38 ± 1.01	3.20 ± 0.32
9	(3E)-3-壬烯醇 (3E)-3-Nonen-1-ol	C₉H₁₈O	-	-	-	-	-	-	1.24 ± 0.07	1.17 ± 0.11
10	2-乙基己醇 2-Ethylhexan-1-ol	C₈H₁₈O	4.28 ± 0.14	1.25 ± 0.08	-	1.93 ± 0.07	-	8.29 ± 0.61	2.08 ± 0.10	2.15 ± 0.09
11	反罗勒烯 Ocimene	C₁₀H₁₆	1.65 ± 0.06	1.53 ± 0.11	2.02 ± 0.08	1.85 ± 0.05	1.55 ± 0.08	-	0.80 ± 0.02	-
12	薄荷烯 cis-p-Menthane	C₁₀H₂₀	3.38 ± 0.04	-	2.03 ± 0.06	-	1.50 ± 0.12	2.48 ± 0.21	-	1.08 ± 0.05
13	柠檬烯 Limonene	C₁₀H₁₆	3.83 ± 0.07	1.42 ± 0.12	1.61 ± 0.12	1.74 ± 0.10	3.29 ± 0.29	1.65 ± 0.15	1.67 ± 0.03	0.94 ± 0.04
14	罗勒烯Ocimene	C₁₀H₁₆	0.18 ± 0.01	3.25 ± 0.24	2.58 ± 0.22	1.44 ± 0.14	1.40 ± 0.10	-	0.94 ± 0.05	-
15	α-蒎烯alpha-Pinene	C₁₀H₁₆	0.07 ± 0.01	2.68 ± 0.16	3.13 ± 0.02	2.52 ± 0.09	5.24 ± 0.34	4.83 ± 0.37	3.60 ± 0.34	2.69 ± 0.27
16	紫苏烯 Perillen	C₁₀H₁₄O	0.34 ± 0.04	0.53 ± 0.05	1.05 ± 0.11	1.60 ± 0.06	1.03 ± 0.02	0.63 ± 0.01	0.54 ± 0.02	-
17	长叶烯 Longifolene	C₁₅H₂₄	1.77 ± 0.13	3.69 ± 0.05	2.05 ± 0.23	0.71 ± 0.03	0.83 ± 0.01	1.46 ± 0.04	1.03 ± 0.02	0.88 ± 0.04
18	丁香烯π Caryophyllen	C₁₅H₂₄	-	-	-	-	-	1.07 ± 0.05	-	-
19	顺-3-乙酸叶醇酯 cis-3-Hexenyl acetate	C₈H₁₄O₂	1.55 ± 0.06	1.61 ± 0.86	2.25 ± 0.61	4.73 ± 1.37	6.11 ± 1.01	2.63 ± 0.31	2.20 ± 0.92	4.09 ± 0.56
20	丙烯酸正己酯 Hexyl acrylate	C₉H₁₆O₂	1.90 ± 0.09	-	-	-	-	2.58 ± 0.18	1.55 ± 0.02	1.08 ± 0.04
21	异丁酸庚酯 Heptyl isobutyrate	C₁₁H₂₂O₂	-	-	0.96 ± 0.03	5.66 ± 0.63	2.45 ± 0.23	6.00 ± 0.93	3.10 ± 0.43	2.47 ± 0.41
22	丁酸庚酯 Heptyl butyrate	C₁₁H₂₂O₂	1.39 ± 0.05	1.31 ± 0.03	4.22 ± 0.08	2.05 ± 0.39	5.02 ± 0.71	4.40 ± 1.21	2.79 ± 0.94	2.18 ± 0.52
23	丁酸丁酯 Butyl butyrate	C₈H₁₆O₂	3.87 ± 0.14	3.07 ± 0.54	5.93 ± 0.35	1.71 ± 0.48	1.25 ± 0.08	-	-	-
	合计		38.89	40.07	48.25	47.75	53.4	52.81	44.83	37.37

表1 波叶金桂’叶片挥发性有机化合物的主要成分(峰面积, A × 105·10 g-1)(平均值±标准误差)

Table 1 The main components of the volatile organic compounds (VOCs) from the leaf of Osmanthus fragrans ‘Boyejingui’ (peak area, A × 105·10 g-1) (mean ± SE)

序号 No.	挥发性有机物 Volatile organic compounds	分子式 Chemical formula	28 ℃				40 ℃
序号 No.	挥发性有机物 Volatile organic compounds	分子式 Chemical formula	对照 Control	轻度干旱 Light drought	中度干旱 Moderate drought	重度干旱 Severe drought	对照 Control	轻度干旱 Light drought	中度干旱 Moderate drought	重度干旱 Severe drought
1	己烯醛 3-Hexenal	C₆H₁₀O	3.33 ± 0.23	0.58 ± 0.65	2.27 ± 0.12	-	3.61 ± 0.26	-	-	1.15 ± 0.05
2	2-甲基-4-戊烯醛 2-Methyl-4-pentenal	C₆H₁₀O	0.99 ± 0.11	2.36 ± 0.23	-	5.97 ± 0.35	-	2.88 ± 0.12	3.14 ± 0.16	1.69 ± 0.13
3	2-已烯醛 (E)-2-Hexenal	C₆H₁₀O	2.48 ± 0.18	2.14 ± 0.23	2.54 ± 0.21	8.67 ± 1.14	4.53 ± 0.54	4.35 ± 0.08	8.01 ± 0.24	7.03 ± 0.19
4	苯基丁醛 4-Phenylbutanal	C₁₀H₁₂O	-	1.85 ± 0.11	-	-	-	5.48 ± 0.13	2.58 ± 0.06	2.74 ± 0.06
5	山梨醛 Sorbaldehyde	C₆H₈O	1.08 ± 0.06	2.82 ± 0.08	1.90 ± 0.04	3.05 ± 0.64	1.80 ± 0.08	-	2.61 ± 0.11	2.33 ± 0.08
6	辛烯醛 2-Ethylhexenal	C₈H₁₄O	-	1.91 ± 0.04	4.86 ± 0.81	-	3.77 ± 0.12	1.37 ± 0.06	0.67 ± 0.02	-
7	壬醛 n-Nonanal	C₉H₁₈O	-	-	-	-	0.76 ± 0.02	1.21 ± 0.04	0.90 ± 0.05	0.53 ± 0.02
8	反-3-己烯醇 trans-3-Hexenol	C₆H₁₂O	6.80 ± 0.45	8.07 ± 0.62	7.85 ± 0.67	4.12 ± 0.35	9.26 ± 1.61	2.50 ± 0.27	5.38 ± 1.01	3.20 ± 0.32
9	(3E)-3-壬烯醇 (3E)-3-Nonen-1-ol	C₉H₁₈O	-	-	-	-	-	-	1.24 ± 0.07	1.17 ± 0.11
10	2-乙基己醇 2-Ethylhexan-1-ol	C₈H₁₈O	4.28 ± 0.14	1.25 ± 0.08	-	1.93 ± 0.07	-	8.29 ± 0.61	2.08 ± 0.10	2.15 ± 0.09
11	反罗勒烯 Ocimene	C₁₀H₁₆	1.65 ± 0.06	1.53 ± 0.11	2.02 ± 0.08	1.85 ± 0.05	1.55 ± 0.08	-	0.80 ± 0.02	-
12	薄荷烯 cis-p-Menthane	C₁₀H₂₀	3.38 ± 0.04	-	2.03 ± 0.06	-	1.50 ± 0.12	2.48 ± 0.21	-	1.08 ± 0.05
13	柠檬烯 Limonene	C₁₀H₁₆	3.83 ± 0.07	1.42 ± 0.12	1.61 ± 0.12	1.74 ± 0.10	3.29 ± 0.29	1.65 ± 0.15	1.67 ± 0.03	0.94 ± 0.04
14	罗勒烯Ocimene	C₁₀H₁₆	0.18 ± 0.01	3.25 ± 0.24	2.58 ± 0.22	1.44 ± 0.14	1.40 ± 0.10	-	0.94 ± 0.05	-
15	α-蒎烯alpha-Pinene	C₁₀H₁₆	0.07 ± 0.01	2.68 ± 0.16	3.13 ± 0.02	2.52 ± 0.09	5.24 ± 0.34	4.83 ± 0.37	3.60 ± 0.34	2.69 ± 0.27
16	紫苏烯 Perillen	C₁₀H₁₄O	0.34 ± 0.04	0.53 ± 0.05	1.05 ± 0.11	1.60 ± 0.06	1.03 ± 0.02	0.63 ± 0.01	0.54 ± 0.02	-
17	长叶烯 Longifolene	C₁₅H₂₄	1.77 ± 0.13	3.69 ± 0.05	2.05 ± 0.23	0.71 ± 0.03	0.83 ± 0.01	1.46 ± 0.04	1.03 ± 0.02	0.88 ± 0.04
18	丁香烯π Caryophyllen	C₁₅H₂₄	-	-	-	-	-	1.07 ± 0.05	-	-
19	顺-3-乙酸叶醇酯 cis-3-Hexenyl acetate	C₈H₁₄O₂	1.55 ± 0.06	1.61 ± 0.86	2.25 ± 0.61	4.73 ± 1.37	6.11 ± 1.01	2.63 ± 0.31	2.20 ± 0.92	4.09 ± 0.56
20	丙烯酸正己酯 Hexyl acrylate	C₉H₁₆O₂	1.90 ± 0.09	-	-	-	-	2.58 ± 0.18	1.55 ± 0.02	1.08 ± 0.04
21	异丁酸庚酯 Heptyl isobutyrate	C₁₁H₂₂O₂	-	-	0.96 ± 0.03	5.66 ± 0.63	2.45 ± 0.23	6.00 ± 0.93	3.10 ± 0.43	2.47 ± 0.41
22	丁酸庚酯 Heptyl butyrate	C₁₁H₂₂O₂	1.39 ± 0.05	1.31 ± 0.03	4.22 ± 0.08	2.05 ± 0.39	5.02 ± 0.71	4.40 ± 1.21	2.79 ± 0.94	2.18 ± 0.52
23	丁酸丁酯 Butyl butyrate	C₈H₁₆O₂	3.87 ± 0.14	3.07 ± 0.54	5.93 ± 0.35	1.71 ± 0.48	1.25 ± 0.08	-	-	-
	合计		38.89	40.07	48.25	47.75	53.4	52.81	44.83	37.37

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