褪黑素对盐碱胁迫下平邑甜茶幼苗生长的影响及其机制

doi:10.16420/j.issn.0513-353x.2022-0642

摘要/Abstract

摘要：

为探究外源褪黑素（melatonin）对盐碱胁迫下平邑甜茶（Malus hupehensis）幼苗生长的影响及其作用机制，以无融合生殖率达到94%的平邑甜茶幼苗为试材，设置3个处理：对照（Hoagland营养液）、盐碱胁迫[100 mmol · L^-1 NaHCO₃︰NaCl = 1︰1（摩尔比）的Hoagland营养液]、盐碱胁迫 + 褪黑素（0.1 mmol · L^-1），15 d后测定幼苗鲜质量、干质量、叶绿素含量、光合速率、矿质元素含量、抗氧化酶活性、渗透物质含量、有机酸含量、内源激素含量以及相关基因的表达量。结果表明，外源褪黑素可以显著降低盐碱胁迫下平邑甜茶幼苗的萎蔫率，并有效提高了盐碱胁迫下植株的生物量和光合效率，同时提高了幼苗中可溶性糖、可溶性蛋白及脯氨酸的含量来缓解渗透胁迫。另外，褪黑素通过增加细胞内有机酸含量和提高AHA家族基因表达量，增强了植株的耐盐碱性;通过调节钠离子转运基因（MhCHX15和MhSOS1）和钾离子转运基因（MhSKOR、MhNHX1、MhNHX2和MhNHX4）的表达，提高细胞质中的钾钠比值以维持离子稳态;并通过提高过氧化物酶（POD）和过氧化氢酶（CAT）活性和调节抗氧化酶基因（MhGPX6、MhpoxN1和MhPER65）的表达，减少盐碱胁迫产生的氧化损伤。此外，外源褪黑素还可通过与赤霉素、生长素、细胞分裂素和茉莉酸甲酯协同作用共同应答盐碱胁迫。综上，盐碱胁迫 + 0.1 mmol · L^-1褪黑素处理通过调节离子平衡、渗透物质、抗氧化酶活性并协同其他内源激素来缓解盐碱胁迫对平邑甜茶幼苗造成的损伤。

关键词: 苹果, 平邑甜茶, 盐碱胁迫, 褪黑素, 高pH胁迫, 渗透平衡, 氧化损伤

Abstract:

To explore the effect of exogenous melatonin on the growth of Malus hupehensis seedlings under saline-alkali stress and its mechanism，M. hupehensis seedlings with 94% apomixis rate were selected as test materials. The control group was irrigated with Hoagland’s nutrient solution（groupⅠ）;The saline-alkali treatment group was irrigated with nutrient solution containing 100 mmol · L^-1 NaHCO₃︰NaCl = 1︰1（groupⅡ）. The group Ⅲ was added with 0.1 mmol · L^-1 melatonin on the basis of groupⅡ. The fresh weight，dry weight，chlorophyll content，photosynthetic rate，mineral element content，antioxidant enzyme activity，osmotic substance content，organic acid content，endogenous hormone content and saline-alkali related gene expression were detected after saline-alkali stress and exogenous melatonin treatment for 15 days. The results showed that application of 0.1 mmol · L^-1 melatonin could significantly enhance the tolerance of M. hupehensis to saline-alkali stress. At first，exogenous melatonin treatment could significantly reduce the wilting rate，and effectively improve the biomass and photosynthesis of M. hupehensis seedlings under salt-alkali stress. Meanwhile，it also increased the content of soluble sugar，soluble protein and proline in seedlings so as to alleviate osmotic stress. In addition，melatonin increased intracellular organic acid content and the expression of AHA family genes to enhanced the saline-alkali tolerance of plants. Moreover，exogenous MT could regulate the expression of Na⁺ transporter genes（MhCHX15 and MhSOS1）and K⁺ transporter genes（MhSKOR，MhNHX1，MhNHX2 and MhNHX4），and enhance the ratio of K⁺/Na⁺ in cytoplasm to maintain ion homeostasis. Melatonin could enhance the activities of POD and CAT and regulate the expressions of antioxidase genes（MhGPX6，MhpoxN1 and MhPER65）to alleviate oxidative damage under saline-alkali stress. Furthermore，exogenous MT could cooperate with gibberellin，auxin，cytokinin and methyl jasmonate responding to saline-alkali stress. In conclusion，exogenous 0.1 mmol · L^-1 melatonin alleviated the damage of saline-alkali stress on M. hupehensis seedlings by regulating ion balance，osmotic substances，antioxidant enzyme activities and coordinating with other endogenous hormones.

Key words: apple, Malus hupehensis, saline-alkali stress, melatonin, high pH stress, osmotic equilibrium, oxidative damage

孙志娟, 刘文杰, 郑晓东, 袭祥利, 马长青, 刘晓丽, 王彩虹, 田义轲. 褪黑素对盐碱胁迫下平邑甜茶幼苗生长的影响及其机制[J]. 园艺学报, 2023, 50(8): 1697-1710.

SUN Zhijuan, LIU Wenjie, ZHENG Xiaodong, XI Xiangli, MA Changqing, LIU Xiaoli, WANG Caihong, TIAN Yike. Effects and Functional Mechanism of Melatonin on the Growth of Malus hupehensis Seedlings Under Saline-Alkali Stress[J]. Acta Horticulturae Sinica, 2023, 50(8): 1697-1710.

图/表 10

表1 qRT-PCR所用的引物序列

Table 1 Primer sequences for qRT-PCR

基因名称 Gene name	上游引物（5′-3′） Forward primer	下游引物（5′-3′） Reverse primer
MhSKOR	CATCCTGACAACTGGTGGTATCG	AATGCCATTTGCCAATAACCT
MhNHX1	TTCTGCGTGAACTTTAGACCCT	AAGACTGAGATTTCCTTTCAAGC
MhNHX2	CCACATTGATTCCAGTATTGCTT	CTCTTGAACTCTCCGTCACATTG
MhNHX4	ACGAAACTCCTTTACTATACAGCCT	TGATACCACAGATAAGTGAGCATAG
MhCHX15	AGATTATCCGTATCCTCTAAGTCTGG	TGGTTCGGCCTATCGTGAAA
MhSOS1	TACACTGTCGCTCTGCTCATCC	CCAGTCGTAAGGGAAAGTGAGC
MhGPX6	TTCCGAGAGTAAATCAATCCACG	AGGCAAACTCTACAATCTCGTCA
MhpoxN1	GCTCCTCCAAATCATTGTTACTG	AAGAAGGACAGAAGCATCACAA C
MhPER65	GGCATTCTATTCCCATTCCCTT	GAGTTGGAAGCGATGAGGAGG
MhAHA1	CCAGAGAAAACAAAAGAGAGTC	TTCACATTCACACCGAGATTG
MhAHA2	AAAGTGTGAAGAAAGAGAAAATGC	AGTTCATTTGCTTGACATTCTTT
MhAHA8	GGTCAAAAGTGTGTTGGTAAGCA	AAACCAAAATCCTCCAACAGC
MhActin	CTTCAATGTGCCTGCCATGTAT	AATTTCCCGTTCAGCAGTAGTG

图1 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株表型的影响

Fig. 1 Effects of exogenous melatonin（MT）on phenotypes of Malus hupehensis plants under（NaCl + NaHCO3）stress

图2 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株萎蔫率及生物量的影响不同小写字母表示差异显著（P < 0.05）。下同。

Fig. 2 Effects of exogenous melatonin（MT）on wilting rate and biomass of Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress Different lowercase letters represent significant differcences（P < 0.05）. The same below.

图3 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株叶片叶绿素含量及光合指标的影响

Fig. 3 Effects of exogenous melatonin（MT）on chlorophyll content and photosynthetic indexes of Malus hupehensis leaves under saline-alkali（NaCl + NaHCO3）stress

图4 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株体内矿质元素含量的影响

Fig. 4 Effects of exogenous melatonin（MT）on the content of mineral elements in Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress

图5 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株活性氧含量及抗氧化酶活性的影响

Fig. 5 Effects of exogenous melatonin（MT）on reactive oxygen content and antioxidant enzyme activities of Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress

图6 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株渗透物质含量的影响

Fig. 6 Effects of exogenous melatonin（MT）on osmotic substance content of Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress

图7 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株有机酸含量的影响

Fig. 7 Effects of exogenous melatonin（MT）on organic acid content of Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress

图8 外源褪黑素（MT）对盐碱（NaCl + NaHCO3）胁迫下平邑甜茶植株体内激素含量的影响

Fig. 8 Effects of exogenous melatonin（MT）on hormone content in Malus hupehensis plants under saline-alkali（NaCl + NaHCO3）stress

图9 外源褪黑素（MT）对平邑甜茶植株盐碱（NaCl + NaHCO3）胁迫应答基因表达的影响

Fig. 9 Effects of exogenous melatonin（MT）on gene expression of salt-alkali（NaCl + NaHCO3）stress response in Malus hupehensis plants

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