外源草酸对采后血橙果实品质和花色苷代谢的影响

doi:10.16420/j.issn.0513-353x.2024-0831

摘要/Abstract

摘要：

为探究草酸处理对采后血橙果实品质和花色苷代谢的影响，以成熟塔罗科血橙（Citrus sinensis L. Osbeck‘Tarocco’）果实为材料，用10 mmol · L^-1草酸浸泡处理5 min，测定贮藏期间果皮色泽、果汁色泽、花色苷、有机酸、糖、总黄酮和总酚含量以及发病率和失水率等指标。同时，还对花色苷生物合成通路相关基因和已报道的参与花色苷代谢调控的重要转录因子进行qRT-PCR分析。结果表明，草酸处理抑制了采后血橙果实贮藏期间花色苷的积累，果皮色泽和果汁色泽较淡，花色苷含量较低；果实柠檬酸含量较对照降低，蔗糖含量明显高于对照。此外，草酸处理明显降低了贮藏后期果实的真菌病害发病率和失重率。草酸处理果实中大多数花色苷合成通路基因（PAL4、CHI和ANS）以及调控基因（NAC、CHR4和PH4）与对照相比在贮藏期间表达显著下调。相关性分析表明，转录因子NAC、WRKY40、Ruby1和PH4的表达趋势与花色苷含量变化趋势显著正相关。研究结果表明草酸处理可对血橙果实采后花色苷的积累产生抑制作用，该抑制作用与花色苷生物合成及调控基因的表达受抑制相关。

关键词: 血橙, 采后, 草酸, 果实品质, 花色苷, 基因表达

Abstract:

To investigate the effects of oxalic acid treatment on the quality and anthocyanin metabolism of postharvest blood orange fruits，mature‘Tarocco’blood orange（Citrus sinensis L. Osbeck‘Tarocco’）fruits were used as materials. The fruits were treated with 10 mmol · L^-1oxalic acid by soaking for 5 min，and indicators such as peel color，juice color，anthocyanin，organic acid，sugar，total flavonoid，and total phenol contents，as well as disease incidence and water loss rate，were measured during storage. Meanwhile，qRT-PCR analysis was performed on genes related to the anthocyanin biosynthesis pathway and reported important transcription factors involved in anthocyanin metabolism regulation. The results showed that oxalic acid treatment inhibited the accumulation of anthocyanin in postharvest blood orange fruits during storage，with lighter peel and juice colors and lower anthocyanin content. The citric acid content in the fruits was lower than that in the control，while the sucrose content was significantly higher than that in the control. In addition，oxalic acid treatment significantly reduced the fungal disease incidence and water loss of fruits in the late storage period. The expression of most genes in the anthocyanin synthesis pathway（PAL4，CHI，and ANS）and regulatory genes（NAC，CHR4，and PH4）in oxalic acid-treated fruits was significantly downregulated compared with the control during storage. Correlation analysis showed that the expression trends of transcription factors NAC，WRKY40，Ruby1，and PH4 were significantly positively correlated with the change trend of anthocyanin content. The research results indicate that oxalic acid treatment can inhibit the accumulation of anthocyanin in postharvest blood orange fruits，and this inhibitory effect is related to the suppression of the expression of genes involved in anthocyanin biosynthesis and regulation.

Key words: blood orange, postharvest, oxalic acid, fruit quality, anthocyanin, gene expression

唐嘉怡, 缪绸雨, 尹豪杰, 吴明月, 曹向敏, 张惠敏, 金燕, 卢晓鹏, 朱亦赤, 李大志, 盛玲. 外源草酸对采后血橙果实品质和花色苷代谢的影响[J]. 园艺学报, 2025, 52(9): 2425-2438.

TANG Jiayi, MIAO Chouyu, YIN Haojie, WU Mingyue, CAO Xiangmin, ZHANG Huimin, JIN Yan, LU Xiaopeng, ZHU Yichi, LI Dazhi, SHENG Ling. The Effect of Exogenous Oxalic Acid Treatment on the Fruit Quality and Anthocyanin Metabolism of Postharvest Blood Orange Fruit[J]. Acta Horticulturae Sinica, 2025, 52(9): 2425-2438.

图/表 9

表1 实时荧光定量分析引物

Table 1 Primers used for qRT-PCR analysis

引物名称 Primer name	正向引物（5′-3′） Forward primer	反向引物（5′-3′） Reverse primer
CsActin	CCAAGCAGCATGAAGATCAA	ATCTGCTGGAAGGTGCTGAG
Cs_ont_8g005310	GTCGTTGAATGCTGCTGAGG	ACGCAAGTCCCTCTTTAGGC
Cs_ont_6g020620	TCTCAGGGAATGGAATGGCG	GTACCATTTAGTCACATCGGCA
Cs_ont_9g012610	TCAAGCGCATGTGTGACAA	GCCATGTACGCACACACATT
Cs_ont_2g025700	AGTTCAAGCGCATGTGTGAC	GCACTTCGACCACCACAATG
Cs_ont_7g003040	GATGCTCTTGTCTCTGCTCCA	GTCCCTCACCGCACTTTCTA
Cs_ont_1g027890	TGGGAGGCACCTTTAAGCTC	TAAGAGAGGCCAGGGGTTCA
Cs_ont_2g002950	TCGAATGGAGGAACCCATCAC	CCATGGTTCACCAACTGAAAGA
Cs_ont_5g038970	CACGTTCGCCAGGAAGAGAT	CAACGCACAAGTTCAACCGT
Cs_ont_3g003380	TCAATGTGCCCACAGAGTTCG	TGGCTCTGCAAGTATCCACAG
Cs_ont_5g040910	CTCCAAGCGACTATACAGAGG	AAGCCAAGTGACAACACTGC
Cs_ont_6g006980	CTTCTTCGACAGCCGTTTGG	GTTTGGCCCTTGGTTTGCAT
Cs_ont_2g004470	GTTTCAGGATCAGGCGGGAA	TTTTCTGGTGCCTGTGGTGT
Cs_ont_2g009870	ATCTGCGACGGCTACAGATG	AGGCGAAAGCTCAGCAAGAT
Cs_ont_2g009920	AAGGGCTGGGACAACTGAAC	TGCAACTGAGAGGGTCCAAC
Cs_ont_2g035300	AACGTCATACCTCTGCTGGC	AAGCGTTCGGTGAATACCCA
Cs_ont_4g001580	GGATTCAACGTGGGTGGACA	TCAGCTTCTTGTTCTCAGTGCT
Cs_ont_4g010710	AACTATGTGGAAGGCCGCTC	ATCGGTGACTTTGAAGCCGT
Cs_ont_5g010100	TTCACCCACCAAAACCAGCT	CTCTTCTCCTTGGTTGGCGT
Cs_ont_6g003620	CCAGCTGTGTCAAGCAGAGA	AACAATGCCGCATGCCATTT
Cs_ont_6g005110	AAGCAGCTGTTCTGTAGGCT	CTGTGGTGCAACCGATGATT
Cs_ont_6g020160	TGCATTTGCCCATCGAGAGT	TTCTGTCGGGGTTTGACGAG

图1 草酸处理对血橙果汁和果皮表型（A）、色差指数CI（B、C）、花色苷含量（D）的影响 * 表示在相同时期草酸与对照在P < 0.05水平的显著性差异，t-test. 下同

Fig. 1 Effects of oxalic acid treatment on the phenotype of juice and peel（A），color difference index CI（B，C）and anthocyanin content（D）of blood oranges * represent significant differences at P < 0.05 between OA and the control at each stage，t-test. The same below

图2 草酸处理对贮藏期间血橙果实糖酸含量的影响

Fig. 2 Effect of OA treatment on content of sugars and organic acids of blood orange fruit during postharvest storage period

图3 草酸处理对血橙果实贮藏期间总黄酮（A）和总酚含量（B）的影响

Fig. 3 Effects of OA treatment on total flavonoid content（A）and total phenol content（B）in blood orange fruit during postharvest storage period

图4 草酸处理对血橙果实贮藏期发病率和失重率的影响

Fig. 4 Effects of OA treatment on incidence rate and weight loss of blood orange fruits during postharvest storage period

图5 草酸处理对血橙果实贮藏期花色苷生物合成相关基因表达的影响

Fig. 5 Effects of OA treatment on expression of genes related to anthocyanin biosynthesis of blood orange fruits during postharvest storage period

图6 草酸处理对血橙果实贮藏期花色苷生物合成调控相关转录因子基因表达的影响

Fig. 6 Effect of OA treatment on the expression of transcription factor genes regulating anthocyanin biosynthesis in blood orange fruits during storage period

表2 10个转录因子表达水平与花色苷含量的相关性

Table 2 Correlation analysis between anthocyanin content and ten transcription factors

元数据ID Mata_ID	基因ID Gene_ID	相关系数 Cor	P值 P value
花色苷Anthocyanin	FAR1(Cs_ont_2g009920)	0.402869	0.219257
花色苷Anthocyanin	HAC12(Cs_ont_6g003620)	0.453251	0.161490
花色苷Anthocyanin	PH4(Cs_ont_2g035300)	0.698900	0.016715
花色苷Anthocyanin	ATC3H64(Cs_ont_4g010710)	0.346240	0.296919
花色苷Anthocyanin	CHR4(Cs_ont_2g009870)	0.153589	0.652082
花色苷Anthocyanin	NAC(Cs_ont_2g004470)	0.882306	0.000324
花色苷Anthocyanin	CH-ZF(Cs_ont_6g020160)	0.340465	0.305587
花色苷Anthocyanin	WRKY40(Cs_ont_4g001580)	0.664538	0.025710
花色苷Anthocyanin	AN1(Cs_ont_5g010100)	0.011326	0.973634
花色苷Anthocyanin	Ruby1(Cs_ont_6g005110)	0.823061	0.001857

图7 与花色苷含量显著相关的转录因子和花色苷合成通路基因的相关网络示意图

Fig. 7 Correlation network diagram among anthocyanin content，transcription factor expression，and anthocyanin biosynthesis pathway gene expression

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