UV-B辐射与低温协同调控荔枝开花的机制研究

doi:10.16420/j.issn.0513-353x.2026-0361

摘要/Abstract

摘要：

荔枝成花需要充足的低温诱导，华南地区出现的“暖冬”常导致荔枝成花障碍。本研究中以5年生‘妃子笑’荔枝盆栽圈枝苗为材料，在人工气候室（15 ℃）中分别设置1.75和3.75 kJ · m^-2· d^-1 UV-B辐射处理，研究UV-B辐射对低温诱导下荔枝生长发育、抗氧化能力和开花的影响。结果表明：高强度UV-B辐射（3.75 kJ · m^-2· d^-1）与低温（15 ℃）结合处理可提高荔枝成花株率和纯花穗比例。低温和UV-B辐射结合可促进荔枝叶片中H₂O₂的积累，但显著降低了叶片的多酚含量和SOD酶活性。叶片中蔗糖和葡萄糖的积累可被短期低温诱导，处理后期均呈降低趋势。低温诱导了LcUVR8-1、LcUVR8-2、LcCOP1、LcCO1、LcCO2、LcFT1和LcFT2基因的表达，UV-B辐射结合低温处理显著上调了LcFT1的表达，但抑制LcFT2的表达。综上，低温结合高强度UV-B辐射处理可能通过促进叶片中H₂O₂的积累，诱导成花基因LcFT1的表达，最终提高荔枝的成花能力。

关键词: 荔枝, UV-B, 低温, 成花, 抗氧化响应, 可溶性糖

Abstract:

Litchi flowering requires sufficient low-temperature induction，but the “warm winters” that frequently occur in southern China often lead to inadequate flowering in litchi. In this study，five-year-old air layered potted‘Feizixiao’litchi plants were placed in a climate chamber at 15 ℃，and two intensities of UV-B radiation（1.75 and 3.75 kJ · m^-2· d^-1，respectively）were applied to investigate the integrative effects of UV-B radiation and low temperature on the growth，development，antioxidant capacity，and flowering of litchi. The results showed that the combination of high-intensity UV-B radiation （3.75 kJ · m^-2· d^-1） and low temperature increased the flowering plant rate and the pure panicle rate in comparison to the control. The combination of UV-B radiation and low temperature enhanced the accumulation of H₂O₂，but significantly reduced the total phenolic content and SOD activity in leaves. The accumulation of sucrose and glucose in leaves was induced by short-term low temperature，but decreased in the late stage of low temperature indcution. Low temperature induced the expression of LcUVR8-1，LcUVR8-2，LcCOP1，LcCO1，LcCO2，LcFT1，and LcFT2；the combination of high-intensity UV-B radiation and low temperature induced the expression of LcFT1 but inhibited the expression of LcFT2. These findings indicate that the accumulation of H₂O₂ by high-intensity UV-B radiation treatment resulting in an up-regulation of LcFT1，may enhance the flowering ability of litchi in the presence of low temperature.

Key words: litchi, UV-B, low temperature, flowering, antioxidant responses, soluble sugars

刘京京, 谢颢阳, 黄晓, 胡郑潇, 曾婉祺, 张惠云, 罗心平, 申济源. UV-B辐射与低温协同调控荔枝开花的机制研究[J]. 园艺学报, 2026, 53(7): 2185-2199.

LIU Jingjing, XIE Haoyang, HUANG Xiao, HU Zhengxiao, ZENG Wanqi, ZHANG Huiyun, LUO Xinping, SHEN Jiyuan. Study on the Integrative Effect of UV-B and Low Temperature on Litchi Flowering Regulatory Mechanism[J]. Acta Horticulturae Sinica, 2026, 53(7): 2185-2199.

图/表 13

图1 人工气候室‘妃子笑’盆栽苗UV-B辐射处理模式

Fig. 1 Treatment scheme for potted ‘Feizixiao’ litchi plants in climate chamber

表1 荔枝基因编码区克隆引物

Table 1 Primers for cloning the coding sequences of litchi genes

基因Gene	正向引物序列（5′-3′）Forward primer sequence	反向引物序列（5′-3′）Reverse primer sequence
LcUVR8-1	CCTTCCATTTTGGACTCTTCTG	GAGCTTCCAGCACCAGTGTAT
LcUVR8-2	CCTCTCTCACGCTCATTCAAC	GATGGGTAAACTGAGTCACTGG
LcCOP1	ATGGAGGTGGCGGAGGTGGAGC	CGTACTTGCGGCTTGAGTCTA
LcCO1	ATGAAAGAAGAGAGCAGCGAC	GGCATTGTTCCTTCATTCTAA

表2 基因qRT-PCR引物

Table 2 Primers for qRT-PCR of target genes

基因Gene	正向引物序列（5′-3′）Forward primer sequence	反向引物序列（5′-3′）Reverse primer sequence
LcUVR8-1	GGGGGTGATGTAAGTGTACCA	GAGCTTCCAGCACCAGTGTAT
LcUVR8-2	CTTGTGGGGGACGTCATACT	GATGGGTAAACTGAGTCACTGG
LcCOP1	CACCAGACATGGATGATACCG	CGTACTTGCGGCTTGAGTCTA
LcCO1	GACCAGACCACGAATCAAGG	ATCACAGACGCCGGATATG
LcCO2	CTCAATACGGGGTCGTTCC	AGCCGACAGATTTCGCAGGAG
LcFT1	GCCGCGGTCTACTTTAATAGC	GCATGTCAAAGCTAGCAAAGA
LcFT2	CTTCAACTGCCAGAGGGAGA	GCATGTCAAAGCTAGCAAAGA
LcActin	ACCGTATGAGCAAGGAAATCACTG	TCGTCGTACTCACCCTTTGAAATC

表3 低温诱导期UV-B辐射处理对‘妃子笑’荔枝开花的影响

Table 3 Effect of UV-B treatment during low-temperature floral induction on flowering of‘Feizixiao’litchi

处理 Treatment	成花株率/% Percentage of flowering plants	成花枝率/% Percentage of flowering branches	纯花穗比例/% Percentage of pure flower spikes	花穗长/cm Inflorescence length	花穗宽/cm Inflorescence width	白点时间/d Days to white spot stage	雌花开放时间/d Days to female flower anthesis	雄花开放时间/d Days to male flower anthesis
CK	50	78.02 ± 6.59 a	42.73 ± 2.73 b	14.23 ± 0.79 a	8.80 ± 0.96 a	7.00 ± 2.00 a	27.00 ± 1.00 a	33.50 ± 0.50 ab
T1	75	66.67 ± 23.13 a	9.52 ± 9.52 c	12.33 ± 1.48 a	6.92 ± 1.14 a	7.00 ± 0.00 a	27.50 ± 2.50 a	32.00 ± 0.00 b
T2	75	93.67 ± 3.19 a	93.33 ± 6.67 a	13.03 ± 0.51 a	8.71 ± 0.45 a	6.25 ± 0.85 a	33.33 ± 4.84 a	35.67 ± 0.67 a

图2 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片光合作用的影响 CK：对照，15 ℃，0.43 kJ · m-2 · d-1；T1：15 ℃，1.75 kJ · m-2 · d-1；T2：15 ℃，3.75 kJ · m-2 · d-1。35 + 7 d为低温处理35 d后升温至25 ℃ 7 d。同期不同字母表示处理组间显著差异（P < 0.05）。下同

Fig. 2 Effect of UV-B treatment during low-temperature floral induction on photosynthesis in‘Feizixiao’litchi leaves CK：The control，15 ℃，0.43 kJ · m-2 · d-1；T1：15 ℃，1.75 kJ · m-2 · d-1；T2：15 ℃，3.75 kJ · m-2 · d-1. 35 + 7 d：25 ℃ for 7 d after 35 d low-temperature treatment. Different letters indicate significant differences between treatments at the 0.05 level. The same below

表4 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片SPAD值的影响

Table 4 Effect of UV-B treatment during low-temperature floral induction on SPAD value of‘Feizixiao’litchi leaves

处理Treatment	0 d	3 d	7 d	21 d	35 d	35 + 7 d
CK	49.75 ± 1.57 a	50.85 ± 1.65 a	50.00 ± 1.29 a	50.58 ± 1.71 a	50.50 ± 1.89 a	50.63 ± 2.02 a
T1	48.20 ± 1.37 a	47.45 ± 1.24 ab	48.60 ± 1.32 a	46.68 ± 1.36 ab	46.65 ± 1.56 a	49.23 ± 1.35 a
T2	45.98 ± 0.26 a	44.93 ± 0.78 b	43.60 ± 1.72 b	45.28 ± 0.56 b	46.60 ± 0.56 a	46.80 ± 0.72 a

图3 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片叶绿素含量的影响 0 d（处理前）样品未设置重复。下同

Fig. 3 Effect of UV-B treatment during low-temperature floral induction on chlorophyll content in‘Feizixiao’litchi leaves There was one biological sample for each treatment on 0 d. The same below

图4 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片中多酚和类黄酮含量的影响

Fig. 4 Effects of UV-B treatment during low-temperature floral induction on total polyphenols and flavonoids content in leaves of ‘Feizixiao’litchi

图5 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片中SOD和POD酶活性的影响

Fig. 5 Effects of UV-B treatment during low-temperature floral induction on SOD and POD activities in leaves of‘Feizixiao’litchi

图6 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片中H2O2含量的影响

Fig. 6 Effect of UV-B treatment during low-temperature floral induction on H2O2 content in leaves of‘Feizixiao’litchi

图7 低温诱导期UV-B辐射处理对‘妃子笑’荔枝叶片可溶性糖含量的影响

Fig. 7 Effects of UV-B treatment during low-temperature floral induction on soluble sugar content in leaves of ‘Feizixiao’ litchi

图8 荔枝中UVR8与其他植物相关基因氨基酸序列比对分析

Fig. 8 Alignment analysis of amino acids sequences of UVR8 between litchi and other related plant species

图9 低温诱导期UV-B辐射处理对叶片中开花相关基因表达量的影响

Fig. 9 Effects of UV-B treatment during low-temperature floral induction on the expression level of flowering-related genes in leaves

参考文献 51

[1]	Abe M, Kobayashi Y, Yamamoto S, Daimon Y, Yamaguchi A, Ikeda Y, Ichinoki H, Notaguchi M, Goto K, Araki T. 2005. FD,a bZIP protein mediating signals from the floral pathway integrator FT at the shoot apex. Science, 309 (5737):1052-1056. doi: 10.1126/science.1115983 URL
[2]	Agati G, Stefano G, Biricolti S, Tattini M. 2009. Mesophyll distribution of‘antioxidant’flavonoid glycosides in Ligustrum vulgare leaves under contrasting sunlight irradiance. Annals of Botany, 104 (5):853-861. doi: 10.1093/aob/mcp177 URL
[3]	Björn L O, Callaghan T V, Johnsen I, Lee J A, Manetas Y, Paul N D, Sonesson M, Wellburn A R, Coop D, Heide-Jørgensen H S, Gehrke C, Gwynn-Jones D, Johanson U, Kyparissis A, Levizou E, Nikolopoulos D, Petropoulou Y, Stephanou M. 1997. The effects of UV-B radiation on European heathland species. Plant Ecology, 128 (3):253-264. doi: 10.1023/A:1009782207376
[4]	Brown B A, Cloix C, Jiang G H, Kaiserli E, Herzyk P, Kliebenstein D J, Jenkins G I. 2005. A UV-B-specific signaling component orchestrates plant UV protection. Proc Natl Acad Sci USA, 102 (50):18225-18230. doi: 10.1073/pnas.0507187102 pmid: 16330762
[5]	Castañeda-Ovando A, Pacheco-Hernández M, Páez-Hernández M, Rodríguez J A. 2009. Chemical studies of anthocyanins:a review. Food Chemistry, 113 (4):859-871. doi: 10.1016/j.foodchem.2008.09.001 URL
[6]	Chauhan S, Chatterjee D, Peer L A, Mir B A, Babbar S B. 2025. Carbon monoxide promotes flowering in Lemna gibba via a nitric oxide-dependent oxidative stress pathway. Planta, 262 (2):47. doi: 10.1007/s00425-025-04775-1
[7]	Chen Tuo, Wang Xunling. 1999. Influence of enhanced UV-B radiation on H₂O₂ metabolism in wheat leaves. Acta Botanica Boreali-Occidentalia Sinica, 19 (2):284-289. (in Chinese)
	陈拓, 王勋陵. 1999. UV-B辐射对小麦叶片H₂O₂代谢的影响. 西北植物学报, 19 (2):284-289.
[8]	Chen Y P. 2009. Response of antioxidant defense system to laser radiation apical meristem of Isatis indigotica seedlings exposed to UV-B. Plant Signaling & Behavior, 4 (7):571-573.
[9]	Choudhary K K, Agrawal S B. 2014. Ultraviolet-B induced changes in morphological,physiological and biochemical parameters of two cultivars of pea(Pisum sativum L.). Ecotoxicology and Environmental Safety,100:178-187.
[10]	Considine M J, Foyer C H. 2014. Redox regulation of plant development. Antioxidants & Redox Signaling, 21 (9):1305-1326.
[11]	Day T A, Demchik S M. 1996. Influence of enhanced UV-B radiation on biomass allocation and pigment concentrations in leaves and reproductive structures of greenhouse-grown Brassica rapa. Vegetatio, 127:109-116. doi: 10.1007/BF00044635 URL
[12]	Ding F, Zhang S W, Chen H B, Su Z X, Zhang R, Xiao Q S, Li H L. 2015. Promoter difference of LcFT1 is a leading cause of natural variation of flowering timing in different litchi cultivars(Litchi chinensis Sonn.). Plant Science,241:128-137.
[13]	Fujibe T, Saji H, Arakawa K, Yabe N, Takeuchi Y, Yamamoto K T. 2004. A methyl viologen-resistant mutant of Arabidopsis,which is allelic to ozone-sensitive rcd1,is tolerant to supplemental ultraviolet-B irradiation. Plant Physiology, 134 (1):275-285. doi: 10.1104/pp.103.033480 URL
[14]	Hao X Y, Sun B H, Song Y X, Zhang J X, Wu J P, Zhang N B, Zhang X M, Yao W K, Xu W R. 2025. Melatonin-mediated physiological and molecular responses to abiotic stress in horticultural crops. Horticultural Plant Journal, 11 (4):1381-1396. doi: 10.1016/j.hpj.2024.08.006 URL
[15]	Hayes F, Williamson J, Mills G. 2012. Ozone pollution affects flower numbers and timing in a simulated BAP priority calcareous grassland community. Environmental Pollution,163:40-47.
[16]	Hideg É, Jansen M A K, Strid Å. 2013. UV-B exposure,ROS,and stress:inseparable companions or loosely linked associates? Trends in Plant Science, 18 (2):107-115. doi: 10.1016/j.tplants.2012.09.003 pmid: 23084465
[17]	Huang Huibai, Chen Houbin. 2003. Anatomical analysis of litchi flower bud differentiation from a phasic view. Journal of Fruit Science, 20 (6):487-492. (in Chinese)
	黄辉白, 陈厚彬. 2003. 以阶段观剖视荔枝的花芽分化. 果树学报, 20 (6):487-492.
[18]	Hong Jiwang, Chen Xingxing, Li Songgang, Zhang Lei, Yang Ziqin. 2015. The impact of H₂O₂ and NO on FT gene expression and floral bud formation in longan. Chinese Journal of Tropical Crops, 36 (11):2002-2006. (in Chinese)
	洪继旺, 陈星星, 李松刚, 张蕾, 杨子琴. 2015. H₂O₂和NO对反季节龙眼成花及FT基因表达的影响. 热带作物学报, 36 (11):2002-2006.
[19]	Kataria S, Jajoo A, Guruprasad K N. 2014. Impact of increasing Ultraviolet-B(UV-B)radiation on photosynthetic processes. Journal of Photochemistry and Photobiology B:Biology,137:55-66.
[20]	Krasensky-Wrzaczek J, Kangasjärvi J. 2018. The role of reactive oxygen species in the integration of temperature and light signals. Journal of Experimental Botany, 69 (14):3347-3358. doi: 10.1093/jxb/ery074 pmid: 29514325
[21]	León-Chan R G, López-Meyer M, Osuna-Enciso T, Sañudo-Barajas J A, Heredia J B, León-Félix J. 2017. Low temperature and ultraviolet-B radiation affect chlorophyll content and induce the accumulation of UV-B-absorbing and antioxidant compounds in bell pepper(Capsicum annuum)plants. Environmental and Experimental Botany,139:143-151.
[22]	Li Fangmin, Chen Yiping, Wang Xunling, Yue Ming. 2006. Combined effects of enhanced UV-B radiation and doubled CO₂ on tomato growth and its fruit quality. Chinese Journal of Applied Ecology, 17 (1):71-74. (in Chinese) pmid: 16689237
	李方民, 陈怡平, 王勋陵, 岳明. 2006. UV-B辐射增强和CO₂浓度倍增的复合作用对番茄生长和果实品质的影响. 应用生态学报, 17 (1):71-74. pmid: 16689237
[23]	Li Jianguo. 2008. The Litchi. Beijing: China Agriculture Press. (in Chinese)
	李建国. 2008. 荔枝学. 北京: 中国农业出版社.
[24]	Liu P, Huang J J, Cai Z Y, Chen H T, Huang X, Yang S N, Su Z X, Azam M, Chen H B, Shen J Y. 2022. Influence of girdling on growth of litchi(Litchi chinensis)roots during cold dependent floral induction. Scientia Horticulturae, 297:110928. doi: 10.1016/j.scienta.2022.110928 URL
[25]	Livak K J, Schmittgen T D. 2001. Analysis of relative gene expression data using real time quantitative PCR and the 2^-ΔΔCT method. Methods,25:402-408.
[26]	Llorens L, Badenes-Pérez F R, Julkunen-Tiitto R, Zidorn C, Fereres A, Jansen M A K. 2015. The role of UV-B radiation in plant sexual reproduction. Perspectives in Plant Ecology, Evolution and Systematics, 17 (3):243-254. doi: 10.1016/j.ppees.2015.03.001 URL
[27]	Lu X Y, Yu S, Lü P T, Chen H B, Zhong S L, Zhou B Y. 2020. Genome-wide transcriptomic analysis reveals a regulatory network of oxidative stress-induced flowering signals produced in litchi leaves. Genes, 11 (3):324. doi: 10.3390/genes11030324 URL
[28]	Mhamdi A, van Breusegem F. 2018. Reactive oxygen species in plant development. Development, 145 (15):dev164376.
[29]	Miller G, Suzuki N, Rizhsky L, Hegie A, Koussevitzky S, Mittler R. 2007. Double mutants deficient in cytosolic and thylakoid ascorbate peroxidase reveal a complex mode of interaction between reactive oxygen species,plant development,and response to abiotic stresses. Plant Physiology, 144 (4):1777-1785. doi: 10.1104/pp.107.101436 pmid: 17556505
[30]	Mittler R, Zandalinas S I, Fichman Y, van Breusegem F. 2022. Reactive oxygen species signalling in plant stress responses. Nature Reviews Molecular Cell Biology, 23 (10):663-679. doi: 10.1038/s41580-022-00499-2
[31]	Musil C F, Wand S J E. 1994. Differential stimulation of an arid-environment winter ephemeral Dimorphotheca pluvialis(L.) Moench by ultraviolet-B radiation under nutrient limitation. Plant, Cell & Environment, 17 (3):245-255. doi: 10.1111/pce.1994.17.issue-3 URL
[32]	Nazir F, Fariduddin Q, Khan T A. 2020. Hydrogen peroxide as a signalling molecule in plants and its crosstalk with other plant growth regulators under heavy metal stress. Chemosphere, 252:126486. doi: 10.1016/j.chemosphere.2020.126486 URL
[33]	Rizzini L, Favory J J, Cloix C, Faggionato D, O‘Hara A, Kaiserli E, Baumeister R, Schäfer E, Nagy F, Jenkins G I, Ulm R. 2011. Perception of UV-B by the Arabidopsis UVR8 protein. Science, 332 (6025):103-106. doi: 10.1126/science.1200660 pmid: 21454788
[34]	Robberecht R, Caldwell M M. 1983. Protective mechanisms and acclimation to solar ultraviolet-B radiation in Oenothera stricta. Plant, Cell & Environment, 6 (6):477-485.
[35]	Saile-Mark M, Tevini M. 1997. Effects of solar UV-B radiation on growth,flowering and yield of central and southern European bush bean cultivars (Phaseolus vulgaris L.). Plant Ecology, 128 (1):115-125. doi: 10.1023/A:1009750612676
[36]	Sami F, Yusuf M, Faizan M, Faraz A, Hayat S. 2016. Role of sugars under abiotic stress. Plant Physiology and Biochemistry,109:54-61.
[37]	She Wenqin, Zhang Yanhui. 2009. Changes of active oxygen metabolism and osmoregulatory substances in litchi leaves during overwintering stage. Chinese Journal of Tropical Crops, 30 (4):490-494. (in Chinese)
	佘文琴, 张延晖. 2009. 越冬期荔枝叶片活性氧代谢和渗透调节物质的变化. 热带作物学报, 30 (4):490-494.
[38]	Shen J Y, Xiao Q S, Qiu H J, Chen C J, Chen H B. 2016. Integrative effect of drought and low temperature on litchi(Litchi chinensis Sonn.)floral initiation revealed by dynamic genome-wide transcriptome analysis. Scientific Reports,6:32005.
[39]	Shi Mingyi, Li Jianli, Zhang Jibin, Mao Qiongli. 2012. Determination of the total flavonoids content in Fagopyrum tartaricum(L.) Gaertn. Pharmacy and Clinics of Chinese Materia Medica, 3 (6):23-25. (in Chinese)
	施明毅, 李建利, 张继斌, 毛琼丽. 2012. 苦荞麦总黄酮含量测定方法研究. 中药与临床, 3 (6):23-25.
[40]	Staxén I, Bornman J F. 1994. A morphological and cytological study of Petunia hybrida exposed to UV-B radiation. Physiologia Plantarum, 91 (4):735-740. doi: 10.1111/ppl.1994.91.issue-4 URL
[41]	Wang M Y, Wang J S, Yang Y H. 2024a. UV-B irradiation enhances the accumulation of beneficial glucosinolates induced by melatonin in Chinese kale sprout. Horticultural Plant Journal, 10 (4):995-1006. doi: 10.1016/j.hpj.2024.05.003 URL
[42]	Wang X Y, Cao M, Li H X, Liu Y, Fan S X, Zhang N, Guo Y D. 2024b. Strategies and prospects for melatonin to alleviate abiotic stress in horticultural plants. Horticultural Plant Journal, 10 (3):601-614. doi: 10.1016/j.hpj.2023.03.011 URL
[43]	Wang Y, Qiu N W, Wang X A, Ma Z Q, Du G Z. 2008. Effects of enhanced UV-B radiation on fitness of an alpine species Cerastium glomeratum Thuill. Journal of Plant Ecology, 1 (3):197-202. doi: 10.1093/jpe/rtn018 URL
[44]	Wen L R, Yang B, Cui C, You L J, Zhao M M. 2012. Ultrasound-assisted extraction of phenolics from longan(Dimocarpus longan Lour.) fruit seed with artificial neural network and their antioxidant activity. Food Analytical Methods,5:1244-1251.
[45]	Xiao Q S, Su Z X, Chen H B, Shen J Y. 2019. Genome-wide identification and involvement of litchi SPL genes in flowering in response to cold and leaf maturity. The Journal of Horticultural Science and Biotechnology, 94 (4):428-440. doi: 10.1080/14620316.2018.1543557 URL
[46]	Yan Bin, Dai Qiujie. 1996. Effects of ultraviolet-B radiation on active oxygen metabolism and membrane system of rice leaves. Acta Phytophysiologica Sinica, 22 (4):373-378. (in Chinese)
	晏斌, 戴秋杰. 1996. 紫外线B对水稻叶组织中活性氧代谢及膜系统的影响. 植物生理学报, 22 (4):373-378.
[47]	Yang Man, An Huaming, Huang Wei, Zhang Xue. 2012. Response of main antioxidant compositions to mechanical wounding and UV-B stress in Rosa roxburghii fruit. Acta Horticulturae Sinica, 39 (4):629-636. (in Chinese)
	杨曼, 安华明, 黄伟, 张雪. 2012. 刺梨果实主要抗氧化组分对机械伤及UV-B胁迫的响应. 园艺学报, 39 (4):629-636.
[48]	Yang Wei. 2010. Effects of UV-B radiation on growth and antioxidant system of wheat seedlings under different temperatures[M. D. Dissertation]. Xi’an: Shaanxi Normal University. (in Chinese)
	杨薇. 2010. UV-B辐射对不同温度下小麦幼苗生长和抗氧化系统的影响[硕士论文]. 西安: 陕西师范大学.
[49]	Yang Z Y, Wang T D, Wang H C, Huang X M, Qin Y H, Hu G B. 2013. Patterns of enzyme activities and gene expressions in sucrose metabolism in relation to sugar accumulation and composition in the aril of Litchi chinensis Sonn. Journal of Plant Physiology,170:731-740.
[50]	Zhou B, Li N, Zhang Z, Huang X, Chen H, Hu Z, Pang X, Liu W, Lu Y. 2012. Hydrogen peroxide and nitric oxide promote reproductive growth in Litchi chinensis. Biologia Plantarum, 56 (2):321-329. doi: 10.1007/s10535-012-0093-3 URL
[51]	Zinser C, Seidlitz H K, Welzl G, Sandermann H, Heller W, Ernst D. 2007. Transcriptional profiling of summer wheat,grown under different realistic UV-B irradiation regimes. Journal of Plant Physiology, 164 (7):913-922. doi: 10.1016/j.jplph.2006.06.006 URL