软枣猕猴桃品种耐热性差异分析与评价

doi:10.16420/j.issn.0513-353x.2023-0676

摘要/Abstract

摘要：

以3个软枣猕猴桃品种为试材，在人工模拟高温与田间自然高温环境下，测定不同品种叶片相对电导率（REC）、气体交换与叶绿素荧光参数、生理生化指标及田间生长指标、热害指数，并采用模糊隶属函数对品种的耐热性进行评价。结果表明：人工模拟高温胁迫下不同品种叶片REC呈“S”形曲线，利用REC配合Logistic方程确定‘龙城2号’‘民大’和‘康甜’高温半致死温度（LT₅₀）分别为44.60、41.92和39.69 ℃，‘龙城2号’和‘民大’在各自LT₅₀下的致死时间分别为85和76 min。温度< 35 ℃对3个品种的REC、叶绿素荧光参数、丙二醛（MDA）含量和游离脯氨酸（Pro）含量无显著影响，> 40 ℃显著影响光合作用与生理代谢，导致细胞膜脂过氧化程度增大，光合效率降低。其中‘龙城2号’的PSⅡ最大光化学效率（F_v/F_m）、PSⅡ实际光化学效率（Φ_PSⅡ）、电子传递速率（ETR）、光化学猝灭系数（q_P）、叶绿素总含量[Chl（a + b）]、净光合速率（P_n）、水分利用效率（WUE）、蒸腾速率（T_r）、Pro均最高且显著高于‘康甜’，而‘康甜’调节性能量耗散量子产量（Y_NPQ）、非调节性能量耗散量子产量（Y_NO）与光合功能限制值（L_PTF）、MDA、REC均最高，其PSⅡ反应中心受高温损伤最重。生长季‘龙城2号’新梢长度、新梢基部直径、叶厚增长较快，叶绿素相对含量（SPAD）较高，田间叶片受害较轻，热害指数最低。LT₅₀与Φ_PSII、ETR、q_P显著正相关，与L_PTF、MDA极显著负相关；田间热害指数与P_n、Chl（a + b）、WUE、F_v/F_m、Φ_PSII、q_P、Pro负相关，与REC、MDA极显著正相关。热害指数与生长速率、LT₅₀均极显著负相关，LT₅₀与生长速率极显著正相关。以半致死温度测定的耐热性与热害指数及以生理指标综合评价结果一致，表明REC、光合参数、生理指标、热害指数可作为软枣猕猴桃耐热性的鉴定指标。综合各项指标确定‘龙城2号’耐热性较强，‘民大’中等，‘康甜’较弱。

关键词: 软枣猕猴桃, 半致死温度, 光合作用, 生理机制, 耐热性

Abstract:

Using three Actinidia arguta cultivars as experimental materials，the relative electrical conductivity（REC），gas exchange and chlorophyll fluorescence parameters，physio-biochemical indexes and field growth performances，heat injury index under artificial-manipulated and ambient high temperature stresses were measured. The heat tolerance was evaluated using fuzzy membership functions over multiple indictor varaibles. The results showed that the relationships between REC and temperature in three tested cultivars conformed well with the Logistic equation. The variation in REC of all A. arguta cultivars with artificial-manipulated increasing temperature followed a S shaped curve. The semi-lethal temperature（LT₅₀）for‘Longcheng 2’‘Minda’and‘Kangtian’were estimated to be 44.60，41.92 and 39.69 ℃，respectively. The failure time of‘Longcheng 2’and‘Minda’under their LT₅₀ were 85 and 76 min，respectively. REC，chlorophyll fluorescence parameters，malondialdehyde（MDA）content and free proline（Pro）content of the three cultivars showed little adjustemnts at < 35 ℃，while photosynthesis and physiological metabolism were drastically mediated at > 40 ℃，resulting in increased membrane peroxidation degree and decreased photosynthetic efficiency. Meanwhile，maximum photochemical efficiency（F_v/F_m），actual photochemical efficiency（Φ_PSⅡ），electron transport rate（ETR），photochemical quenching（q_P），total chlorophyll content[Chl（a + b）]，net photosynthetic rate（P_n），water use efficiency （WUE），transpiration rate（T_r）and Pro of‘Longcheng 2’were the highest among three cultivars，and significantly higher than those of‘Kangtian’. On the contrary，‘Kangtian’was the highest with respect to the yield for dissipation by down-regulation（Y_NPQ），the yield of other non-photochemical losses（Y_NO），the relative limitation of photosynthesis（L_PTF），MDA and REC，indicating that its’ PSⅡ reaction center had been most damaged by high temperature. In the growing season，field-grown‘Longcheng 2’had faster growth in shoot length，base diameter，leaf thickness and higher relative chlorophyll content（SPAD），and its growth rate was significantly higher than that of the other two cultivars，with slighter leaf damage and the lowest heat injury index under high temperature. LT₅₀ was positively correlated with Φ_PSⅡ，ETR and q_P（P < 0.05），and negatively correlated with L_PTF and MDA（P < 0.01）. Heat injury index was negatively correlated with P_n，Chl（a + b），WUE，F_v/F_m，Pro，Φ_PSⅡ and q_P（P < 0.01），but positively correlated with REC and MDA（P < 0.01）. Simutenously，the heat injury index was negatively correlated with growth rate and LT₅₀（P < 0.01），and the LT₅₀ was positively correlated with the growth rate（P < 0.01）. The rank of heat tolerance in terms of semi-lethal temperature was consistent with the results of a comprehensive evaluation based on physiological indexes and heat injury index. These results indicate that REC，photosynthetic parameters，physio-biochemical indexes and heat injury index can be used to indicate heat tolerance of A. arguta cultivars. Altogether，the rank of heat tolerance was determined as follow：‘Longcheng 2’（strong heat tolerance）>‘Minda’（medium heat tolerance）>‘Kangtian’（weak heat tolerance）.

Key words: Actinidia arguta, semi-lethal temperature, photosynthesis, physiological mechanism, heat tolerance

王海珍, 应瑶琳, 王雨晴, 吕瑞恒, 韩路. 软枣猕猴桃品种耐热性差异分析与评价[J]. 园艺学报, 2024, 51(12): 2857-2870.

WANG Haizhen, YING Yaolin, WANG Yuqing, LÜ Ruiheng, HAN Lu. Analysis and Evaluation of Heat Tolerance of Actinidia arguta Cultivars in Extreme Arid Area[J]. Acta Horticulturae Sinica, 2024, 51(12): 2857-2870.

图/表 11

图1 不同品种离体叶片在不同温度下1 h时的相对电导率不同小写字母表示品种间差异显著（P < 0.05）。

Fig. 1 Relative electric conductivity of leaves of different cultivars at different temperatures for 1 h Different lowercase letters indicate significant differences among different cultivars at 0.05 level.

表1 Logistic拟合方程及半致死温度

Table 1 The parameters of Logistic fitting equation and semi-lethal temperature

品种 Cultivar	回归方程 Regression equation	拟合度 R²	半致死温度/℃ LT₅₀	排序 Ranking
龙城2号Longcheng 2	y = 100/（1 + 45.696e^0.0857^t）	0.9674^**	44.60	1
民大Minda	y = 100/（1 + 45.332e^0.0910^t）	0.9439^**	41.92	2
康甜Kangtian	y = 100/（1 + 47.948e^0.0975^t）	0.9583^**	39.69	3

图2 不同品种离体叶片在半致死温度下不同时间的相对电导率不同小写字母表示品种间差异显著（P < 0.05）。

Fig. 2 Relative electric conductivity of leaves of different cultivars at semi-lethal temperature with different times Different lowercase letters indicate significant differences among different cultivars at 0.05 level.

图3 不同软枣猕猴桃品种叶绿素荧光参数对高温的响应不同小写字母表示品种间差异显著（P < 0.05）。

Fig. 3 The high temperature responses of chlorophyll fluorescence parameters of different Actinidia arguta cultivars Different lowercase letters indicate significant difference at P < 0.05 among different cultivars.

图4 高温对不同软枣猕猴桃品种叶片脯氨酸（Pro）和丙二醛（MDA）含量的影响不同小写字母表示相同温度品种间差异显著（P < 0.05）。

Fig. 4 The effect of high temperature on free proline（Pro）and malondialdehyde（MDA）content in leaves with different cultivars Different lowercase letters indicate significant differences among different cultivars in the same temperature（P < 0.05）.

表2 田间自然高温下不同软枣猕猴桃品种的枝叶生长和叶片叶绿素含量增长速率

Table 2 The effect of natural high temperature on growth rates of shoot-leaf and chlorophyll content with different cultivars in field

月份 Month	品种 Cultivar	新梢长/（cm · d^-1） Shoot length	新梢基径/（mm · d^-1） Base diameter	叶厚/（mm · d^-1） Leaf thickness	叶绿素含量（SPAD · d^-1） Relative chlorophyll content
4月 April	龙城2号Longcheng 2	3.459 ± 0.178 a	0.118 ± 0.006 a	0.0022 ± 0.0002 a	0.554 ± 0.029 b
	民大Minda	2.299 ± 0.143 b	0.113 ± 0.004 a	0.0022 ± 0.0002 a	0.676 ± 0.042 a
	康甜Kangtian	1.644 ± 0.087 c	0.095 ± 0.004 b	0.0019 ± 0.0001 a	0.327 ± 0.018 c
5月 May	龙城2号Longcheng 2	3.120 ± 0.161 a	0.048 ± 0.003 a	0.0020 ± 0.0002 a	0.342 ± 0.020 a
	民大Minda	1.151 ± 0.008 b	0.045 ± 0.003 a	0.0017 ± 0.0001 a	0.321 ± 0.016 a
	康甜Kangtian	1.097 ± 0.052 b	0.033 ± 0.002 b	0.0015 ± 0.0001 a	0.281 ± 0.012 a
6月 June	龙城2号Longcheng 2	0.432 ± 0.022 a	0.034 ± 0.002 a	0.0018 ± 0.0002 a	0.209 ± 0.011 a
	民大Minda	0.280 ± 0.014 b	0.030 ± 0.001 a	0.0015 ± 0.0001 a	0.155 ± 0.008 ab
	康甜Kangtian	0.109 ± 0.006 c	0.019 ± 0.001 b	0.0015 ± 0.0001 a	0.097 ± 0.005 b

表3 不同软枣猕猴桃品种叶绿素荧光参数对田间自然高温的响应

Table 3 High temperature response of chlorophyll fluorescence parameters of different Actinidia arguta cultivars in field

品种Cultivar	F_v/F_m	F_v′/F_m′	Φ_PSⅡ	q_P	NPQ	Y_NPQ	Y_NO	L_PTF
龙城2号 Longcheng 2	0.74 ± 0.03 a	0.47 ± 0.02 a	0.20 ± 0.01 a	0.41 ± 0.02 a	2.31 ± 0.11 b	0.60 ± 0.02 ab	0.21 ± 0.01 b	0.43 ± 0.02 c
民大Minda	0.71 ± 0.02 a	0.41 ± 0.01 b	0.18 ± 0.01 ab	0.38 ± 0.02 a	2.27 ± 0.10 b	0.58 ± 0.01 b	0.25 ± 0.01 ab	0.53 ± 0.02 b
康甜Kangtian	0.64 ± 0.01 b	0.39 ± 0.02 b	0.13 ± 0.01 b	0.29 ± 0.01 b	2.89 ± 0.13 a	0.61 ± 0.01 a	0.26 ± 0.01 a	0.59 ± 0.03 a

图5 田间自然高温下不同品种的生理指标不同小写字母表示品种之间差异显著（P < 0.05）；* 表示同一品种不同时间差异显著。

Fig. 5 The effect of natural high temperature on physiological indexes of different Actinidia arguta cultivars in field Different lowercase letters indicate significant differences among different cultivars（P < 0.05），and * indicate significant differences（P < 0.05）among different time in the same cultivar.

表4 半致死温度（LT50）、热害指数（HII）与生长速率（AGR）、生理指标的相关系数

Table 4 Correlation coefficient between growth，physiological indexes and semi-lethal temperature，heat injury index

指标Index	LT₅₀	HII	指标Index	LT₅₀	HII
F_v/F_m	0.8627^*	-0.9698^**	AGR	0.9360^**	-0.9588^**
Φ_PSII	0.9591^**	-0.9554^*	HII	-0.9298^**
ETR	0.9499^**		P_n		-0.9694^**
q_P	0.9286^**	-0.9365^**	Chl（a + b）		-0.9274^**
q_N	-0.9285^**		Car		-0.9396^**
L_PTF	-0.9668^**		REC		0.9934^**
MDA	-0.9544^**	0.9795^**	WUE		-0.9219^**
Pro	-0.7421	-0.9499^**

表5 软枣猕猴桃光合速率与环境因子的逐步回归分析

Table 5 Step regression analysis of photosynthetic rate and environmental factors in Actinidia arguta cultivars

指标Index	B	SE	Beta	t	VIF
常数 Constant	5.1000	0.3580		14.254
空气温度 Air temperature	-0.0910	0.0120	-0.326	-7.938	2.841
饱和水汽压差 Vapor pressure deficit	-0.1300	0.0290	-0.196	-4.425	3.302
光合有效辐射 Photo synthetically active radiation	0.0001	0.0001	0.252	8.657	1.420

表6 不同品种各生理指标隶属函数值与综合评价结果

Table 6 Subordinator functional components and the integrated evaluation index of physiology parameters for different cultivars

品种 Cultivar	隶属函数值 Subordinative function value															CE
品种 Cultivar	SL	LT	F_v/F_m	Φ_PSII	q_P	NPQ	L_PTF	P_n	T_r	WUE	Chl (a + b)	Car	REC	MDA	Pro	CE
龙城2号 Longcheng 2	1.000	1.000	1.000	1.000	1.000	0.939	1.000	1.000	1.000	1.000	1.000	1.000	1.000	1.000	1.000	0.996
民大Minda	0.207	0.445	0.772	0.463	0.712	1.000	0.380	0.566	0.075	0.990	0.723	0.817	0.906	0.760	0.800	0.642
康甜Kangtian	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000	0.000

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