从调控蔗糖代谢的角度分析了外源果糖处理减轻雷竹笋(Phyllostachys praecox f. Preveynalis)采后木质化败坏症状的机理。将雷竹笋分别用0(对照)、10、20、30或40 mmol/L的果糖溶液进行喷淋处理(20 ℃),随后置于4 ℃、相对湿度80%~90%条件下冷藏20 d,期间每隔5 d取样测定笋体木质化和品质参数、可溶性糖和尿苷二磷酸葡萄糖(uridine diphosphate glucose,UDPG)含量以及蔗糖代谢和苯丙烷类代谢途径相关基因的表达丰度。结果表明,20 mmol/L果糖处理较其他处理能更为显著地延缓雷竹笋冷藏期间硬度、失重率、笋体褐变度以及木质素和纤维素含量的上升,提高出汁率,维持品质和抗氧化活性,从而有效控制笋体冷藏期间木质化败坏症状的发展。20 mmol/L果糖处理可显著上调雷竹笋在冷藏期间蔗糖合成方向的基因表达量(PpSS2、PpSPS和PpSPP),促进蔗糖合成并减少UDPG的积累;同时,经果糖处理的雷竹笋中苯丙烷类代谢途径相关基因(PpPAL、Pp4CL、PpCAD、PpF5H、PpCCOMT和PpPOD)的表达丰度显著低于对照水平,并伴随着木质素合成量的显著下降。由此推测,果糖处理可通过调控雷竹笋蔗糖代谢途径的方式提高雷竹笋冷藏期间蔗糖合成量,减少木质素合成中间体UDPG的积累,从而竞争性抑制笋体以苯丙烷类代谢途径合成木质素,最终抑制笋体木质化的发生。
The present study aimed to analyze the effects of exogenous fructose treatment on alleviating lignification deterioration of bamboo shoots (Phyllostachys praecox) during cold storage and the involved mechanism on basis of the regulation of sucrose metabolism. The bamboo shoots were sprayed by the fructose solutions at different concentrations (0,10,20,30 or 40 mmol/L) at 20 ℃ and air-dried respectively. Then the bamboo shoots were stored at 4 ℃ for 20 days. During the storage,the bamboo shoots were collected at a 5-day interval for measurement of the lignification,quality parameters,contents of soluble sugars,uridine diphosphate glucose (UDPG) as well as the expressions of a set of relevant genes involved in sucrose and phenylpropanoid metabolism pathway. The results exhibited that the treatment of bamboo shoots with 20 mmol/L fructose could delay the increase of firmness, the loss of weight,browning degree and the contents of lignin and cellulose. And,it could enhance the extractable juice rate and maintain the overall quality and antioxidant activity,thus contributing to the alleviation of lignification deterioration during the cold storage. Meanwhile,the fructose treatment significantly up-regulated the expression levels of PpSS2,PpSPS and PpSPP which could promote the sucrose synthesis and decrease the accumulation of UDPG in bamboo shoots during the storage. On the contrary,the expression levels of PpPAL,Pp4CL,PpCAD,PpF5H,PpCCOMT and PpPOD related to phenylpropanoid metabolism in fructose-treated bamboo shoots were down-regulated remarkably,accompanied with the clear decrease in lignin content. Therefore,it is deduced that the fructose treatment could enhance sucrose synthesis and decrease UDPG accumulation in terms of the substances for lignin synthesis,resulting in competitive inhibition of phenylpropanoid pathway for lignin synthesis and finally inhibiting the lignification deterioration in bamboo shoots during cold storage.
[1] JIANG P K,XU Q F,XU Z H,et al.Seasonal changes in soil labile organic carbon pools within a Phyllostachys praecox stand under high rate fertilization and winter mulch in subtropical China[J]. Forest Ecology and Management,2006,236(1): 30-36.
[2] CHEN H,LING J,WU F,et al.Effect of hypobaric storage on flesh lignification,active oxygen metabolism and related enzyme activities in bamboo shoots[J]. LWT-Food Science and Technology,2013,51(1): 190-195.
[3] LUO Z,XU X,YAN B.Use of 1-methylcyclopropene for alleviating chilling injury and lignification of bamboo shoot (Phyllostachys praecox f. prevernalis) during cold storage[J]. Journal of the Science of Food and Agriculture,2008,88(1): 151-157.
[4] 黄劲松,高云,郭军,等. 采后竹笋木质化研究进展[J]. 广东农业科学,2018,45(7): 105-110.
[5] VOGT T.Phenylpropanoid biosynthesis[J]. Molecular Plant,2010,3(1): 2-20.
[6] WANG K,JIN P,SHANG H,et al.Effect of methyl jasmonate in combination with ethanol treatment on postharvest decay and antioxidant capacity in Chinese bayberries[J]. Journal of Agricultural and Food Chemistry,2010,58(17): 9 597-9 604.
[7] 李园园,王莉,周梦洁,等. 2,4-表油菜素内酯对草莓果实贮藏品质及抗氧化活性的影响[J]. 食品科学,2018,39(1): 279-284.
[8] EL-METWALLY M A,TARABIH M E,EL-ERYAN E E. Effect of application of β-aminobutyric acid on maintaining quality of crimson seedless grape and controlling postharvest diseases under cold storage condition[J]. Plant Pathology Journal,2014,3(3): 139-151.
[9] 汪开拓,郑永华. 硼处理对杨梅果实采后贮藏期间蔗糖代谢及花色苷合成的影响[J]. 食品与发酵工业,2012,38(9): 137-143.
[10] WANG K,LIAO Y,CAO S,et al.Effects of benzothiadiazole on disease resistance and soluble sugar accumulation in grape berries and its possible cellular mechanisms involved[J]. Postharvest Biology and Technology,2015,102(1): 51-60.
[11] FERRER J L,AUSTIN M B,STEWART C,et al.Structure and function of enzymes involved in the biosynthesis of phenylpropanoids[J]. Plant Physiology and Biochemistry,2008,46(3): 356-370.
[12] XU F,TANG Y,DONG S,et al.Reducing yellowing and enhancing antioxidant capacity of broccoli in storage by sucrose treatment[J]. Postharvest Biology and Technology,2016,112(2): 39-45.
[13] 汪开拓,唐利,李静,等. 茉莉酸甲酯处理对冷藏雷竹笋内源激素水平及木质素合成相关酶活性的影响[J]. 中国蔬菜,2012,1(2): 54-60.
[14] CAO S,ZHENG Y,WANG K,et al.Effect of methyl jasmonate on cell wall modification of loquat fruit in relation to chilling injury after harvest[J]. Food Chemistry,2010,118(3): 641-647.
[15] JIANG J,JIANG L,LUO H B,et al.Establishment of a statistical model for browning of fresh-cut lotus root during storage[J]. Postharvest Biology and Technology,2014,92(3): 164-171.
[16] 杨光,王丹丹,李琴,等. 不同贮藏温度对雷竹笋品质的影响[J]. 食品与发酵工业,2017,43(5): 233-239.
[17] KAMPFENKEL K,VANMONTAGU M,INZE D.Extraction and determination of ascorbate and dehydroascorbate from plant tissue[J]. Analytical Biochemistry,1995,225(1): 165-167.
[18] RUFINO M S M,FERNANDES F A N,ALVES R E,et al. Free radical-scavenging behaviour of some north-east Brazilian fruits in a DPPH system[J]. Food Chemistry,2009,114(2): 693-695.
[19] WILSON A M,WORK T M,BUSHWAY A A,et al.HPLC determination of fructose,glucose,and sucrose in potatoes[J]. Journal of Food Science,1981,46(1): 300-301.
[20] GOULARD F,DIOURIS M,DESLANDES E,et al.An HPLC method for the assay of UDP-glucose pyrophosphorylase and UDP-glucose-4-epimerase in Solieria chordalis(Rhodophyceae)[J]. Phytochemical Analysis,2001,12(6): 363-365.
[21] LIVAK K J,SCHMITTGEN T D.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method[J]. Methods,2001,25(4): 402-408.
[22] LASTDRAGER J,HANSON J,SMEEKENS S.Sugar signals and the control of plant growth and development[J]. Journal of Experimental Botany,2014,65(3): 799-807.
[23] CORDENUNSI B R,OLIVEIRA DO NASCIMENTO J R,GENOVESE M I,et al. Influence of cultivar on quality parameters and chemical composition of strawberry fruits grown in Brazil[J]. Journal of Agricultural and Food Chemistry,2002,50(9): 2 581-2 586.
[24] HAN A,CAO S,LI Y,et al.Sucrose treatment suppresses programmed cell death in broccoli florets by improving mitochondrial physiological properties[J]. Postharvest Biology and Technology,2019,156(1): 110 932-110 938.
[25] 汤月昌,许凤,王鸿飞,等. 葡萄糖处理对青花菜品质和抗氧化性的影响[J]. 食品科学,2014,35(14): 205-209.
[26] 杨少华,王丽,穆春,等. 蔗糖调节拟南芥花青素的生物合成[J]. 中国生物化学与分子生物学报,2011,27(4): 364-369.
[27] 王春幸,王鸿飞,熊茜,等. 果糖处理对费菜总黄酮及其相关酶的影响[J]. 食品工业科技,2017,38(5): 124-129.
[28] BLANCO-HERRERA F,MORENO A A,TAPIA R,et al.The UDP-glucose: glycoprotein glucosyltransferase (UGGT),a key enzyme in ER quality control,plays a significant role in plant growth as well as biotic and abiotic stress in Arabidopsis thaliana[J]. BMC Plant Biology,2015,15(1): 127-138.
[29] CIERESZKO I,JOHANSSON H,KLECZKOWSKI L A.Sucrose and light regulation of a cold-inducible UDP-glucose pyrophosphorylase gene via a hexokinase-independent and abscisic acid-insensitive pathway in Arabidopsis[J]. Biochemical Journal,2001,354(1): 67-72.
[30] WANG K,WU D,GUO D,et al.β-aminobutyric acid induces disease resistance against Botrytis cinerea in grape berries by a cellular priming mechanism[J]. Acta Alimentaria,2019,48(2): 177-186.