采后处理对延缓青花菜衰老的作用及其机理研究
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摘要
青花菜(Brassica oleracea var. italica)因含有丰富的Vc、硫代葡萄糖苷和酚类等营养物质而备受消费者青睐。但青花菜在未成熟时采收,采后在室温下花蕾迅速开放,花球衰老变黄,从而影响外观品质,降低商品价值。因此,研究青花菜采后衰老黄化机理及其控制技术,已成为解决青花菜贮运保鲜问题的关键。本论文以青花菜为试材,在研究不同贮藏温度下青花菜颜色变化和衰老黄化规律的基础上,重点研究了6-苄基腺嘌呤(6-BA)、1-甲基环丙烯(1-MCP)和乙醇处理对延缓青花菜采后黄化的作用及其机理,并探索了6-BA和1-MCP复合处理对延缓青花菜衰老的效果。此外,还研究了1-MCP处理对青花菜采后糖代谢及其相关酶的调控,为糖作为信号分子调控青花菜的衰老提供依据。研究结果分述如下：
(1)研究了不同贮藏温度(273、278、283、293和303K)对青花菜在贮藏期间颜色和叶绿素含量的影响。结果表明,在283、293和303K贮藏时,青花菜-a／b值和叶绿素含量随着贮藏时间延长逐渐下降,温度越高,下降越快。而在273和278K贮藏时,-a／b值和叶绿素含量无显著变化,说明低温能有效延缓青花菜黄化和叶绿素降解。基于Arrhenius动力学方程分别建立了青花菜-a/b值、叶绿素含量与贮藏时间和温度之间的动力学模型,再根据-a/b值与叶绿素含量变化的线性关系,进一步建立了基于-a/b值的叶绿素含量预测模型。该预测模型较好的预测在273-305K温度下青花菜叶绿素含量的变化状况,为应用物理颜色参数检测方法来检测青花菜品质提供依据。
(2)研究了6-BA处理对青花菜颜色、抗氧化能力、总酚、总硫代葡萄糖苷和萝卜硫素含量的影响。结果发现200mg/L6-BA处理显著抑制了L*值和丙二醛(MDA)的增长,以及H值的下降。6-BA处理通过调控叶绿素酶和脱镁螯合酶活性来抑制叶绿素降解。与对照组相比,6-BA处理增强了DPPH自由基清除能力,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性,减弱了过氧化物酶(POD)活性。6-BA处理显著增加了总酚、总硫代葡萄糖苷和萝卜硫素含量。这些结果说明,6-BA处理能保持青花菜品质,延缓衰老和改善其营养保健价值。
(3)研究了乙醇处理对青花菜表观颜色、抗氧化酶活性和活性成分的影响。500μl/l乙醇处理显著地延长了青花菜货架寿命,抑制了L*值的上升和H值的下降。通过抑制叶绿素酶和脱镁螯合酶活性来延缓叶绿素降解。乙醇处理增强了DPPH自由基清除能力、SOD、APX和CAT活性。乙醇处理提高了总酚、总硫代葡萄糖苷和萝卜硫素含量。这些结果说明应用乙醇能延长青花菜货架期,维持感官质量和营养品质。
(4)2.5μl/11-MCP处理能有效地延长青花菜的货架寿命,抑制颜色参数H值的下降和L*值的上升,通过抑制叶绿素酶、脱镁螯合酶和POD的活性来延缓叶绿素的降解,维持其表观绿色。1-MCP处理提高了总酚含量和SOD、APX和CAT等抗氧化酶活性,使得抗氧化能力增强。同时,也延缓了总硫代葡萄糖苷和萝卜硫素含量下降,改善了青花菜的营养保健功能。
(5)离体组织培养青花菜花蕾实验结果显示,不同种类可溶性糖能抑制叶绿素的降解。研究了1-MCP处理对采后青花菜叶绿素含量、糖代谢及其相关酶活性的影响。结果表明,与对照组相比,2.5μl/l1-MCP处理组青花菜维持更高的叶绿素a和b含量。与对照组相比,1-MCP处理增强了蔗糖磷酸合酶(SPS)和蔗糖合酶合成方向(SS-S)活性,抑制了蔗糖合酶分解方向(SS-C)、葡萄糖激酶(GK)和UDP-葡萄糖焦磷酸化酶(UGPaSe)活性。1-MCP减少了蔗糖、果糖和葡萄糖含量下降,维持了较高水平糖含量。这些结果说明,糖在青花菜衰老过程中起重要作用,1-MCP处理通过调控糖代谢和维持较高水平可溶性糖来延缓青花菜衰老。
(6)研究了2.5μl/l1-MCP和200mg/L6-BA复合处理对15℃贮藏条件下青花菜表观颜色、抗氧化酶和保健成分(萝卜硫素和硫代葡萄糖苷)的影响。复合处理比单一处理更显著延缓了青花菜贮藏期间衰老的发生并维持了花蕾的品质。1-MCP复合6-BA处理保持高水平的H值和叶绿素含量,抑制了L*值的增加和POD活性,提高了CAT、APX和SOD的活性。该复合处理较单一处理更为有效地延缓硫代葡萄糖苷的分解和诱导了青花菜中萝卜硫素的形成,维持了青花菜的营养品质。
(7)6-BA、1-MCP和乙醇处理能抑制青花菜叶绿素降解,与对照组相比,6-BA处理对BocHL1、BocHL2和BocHL3的表达没有显著影响,乙醇处理抑制了BocHL2和BoCHL3的表达,而1-MCP处理却诱导了BOCHL1、BocHL2和BoCHL3的表达,这暗示着BoCHL1、BoCHL2和BoCHL3可能在叶绿素降解过程中并没有起到关键的作用。乙醇处理延缓了青花菜衰老时BoPAO正调控,1-MCP则诱导了BoPAO的表达。这些结果表明,不同采后处理通过对叶绿素降解酶基因的调控作用,从而抑制了青花菜的黄化。
Broccoli (Brassica oleracea L. ssp. italica) is known to have high nutritional value with significant contents of vitamins, glucosinolates and phenolic compounds. The inflorescences are harvested while the floral heads, branchlets, and are totally immature, with the sepals completely surrounding the flower. However, broccoli deteriorates very quickly after harvest at ambient temperature due to the loss of green color and the consequent yellowing of florets, which decreases the commercial acceptance of the broccoli florets. Therefore, it is crucial to study the regulation and mechanism of quality deterioration of broccoli florets after harvest for transportation and storage. Basing on the change of color and yellowing of broccoli florets upon various temperatures storage, the present study was designed to investigate the effect of6-benzylaminopurine (6-BA),1-methylcyclopropene (1-MCP) and ethanol on inhibiting yellowing of postharvest broccoli florets and the possible mechanisms involved. The effect of1-MCP in combination with6-BA in delaying senescence on broccoli florets was also assessed. Furthermore, the effect of1-MCP treatment on regulation sugar metabolism and involvement of enzymes in broccoli florets was investigated, which laid a theoretical basis for sugars acting as signaling molecules in regulating senescence of broccoli florets. The results were as follows:
(1) The effect of different storage temperatures (273,278,283,293and303K) on color and chlorophyll content of postharvest broccoli florets was investigated. The results showed that-a/b value and chlorophyll content of broccoli florets markedly decreased during stored at283,293and303K. The increase of storage temperature, change scope of-a/b value and chlorophyll content was also promoted. There were no significant differences in color and chlorophyll content of broccoli florets during stored at273and278K, which suggested that low temperature can delay the yellowing and chlorophyll degradation of broccoli. The kinetic models of-a/b value and chlorophyll content with respect to storage time and temperature were established based on the Arrhenius equation, respectively. Then, a prediction model for chlorophyll content based on-a/b value was developed according to the linear relationship between chlorophyll content and-a/b value. Finally, the chlorophyll content of broccoli can be accurately predicted at the storage temperature from273to305K by the prediction model, which confirms the feasibility of applying the objective color parameter measurement for assessment of broccoli quality.
(2) The effect of6-BA on color, antioxidant activity, contents of total phenols, glucosinolate and sulforaphane in broccoli florets was investigated. The results showed that200mg/L6-BA treatment markedly inhibited the increase of L value and malondialdehyde (MDA) content, and retarded the decrease of H value.6-BA treatment reduced rate of chlorophyll degradation by regulating the activities of chlorophyllase and Mg-dechelatase. When compared to control florets, the1,1-diphenyl-2-picrylhydrazyl (DPPH) radical scavenging activity and the activities of superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) were enhanced in florets treated with6-BA, whereas the activity of peroxidase (POD) was reduced. The contents of total phenols, glucosinolate and sulforaphane were increased after treated with6-BA. These results indicated that6-BA could maintain quality, delay senescence and improve the nutritional value of broccoli.
(3) The effect of cthanol treatment on visual quality, antioxidant enzymes and bioactive compounds in broccoli florets was investigated.500μ1/1ethanol treatment markedly extended shelf life, inhibited the increase of L*value and the decrease of H value. It reduced the rate of chlorophyll degradation by inhibiting the activities of chlorophyllase and Mg-dechelatase. In addition, the DPPH radical scavenging activity and the activities of SOD, APX and CAT were enhanced in florets treated with ethanol. The contents of total phenolic compounds, glucosinolates and sulforaphane were also profoundly increased after treated with ethanol. These results indicated that a application of ethanol can prolong shelf life, maintain visual quality and the nutritional value of broccoli florets.
(4)2.5μl/11-MCP treatment significantly extended shelf life, inhibited the increase of L value and the decrease of H value.1-MCP might delay chlorophyll degradation and florets yellowing by inhibiting the activities of chlorophyllase, Mg-dechelatase and POD. The enhancement in antioxidant activity by1-MCP may be due to increased phenolic content and higher activities of antioxidant enzymes including CAT, SOD and APX. Thus1-MCP has a potential application in postharvest treatment for prolonging shelf life, maintaining quality and improving the health benefit of broccoli florets.
(5) In vitro incubation of broccoli florets showed that various soluble sugars can retard the chlorophyll degradation in broccoli florets. The effect of1-methylcyclopene (1-MCP) on chlorophyll content, sugar metabolism and involvement of enzymes in broccoli florets was investigated. The results showed that1-MCP treatment maintained higher chlorophyll a and b contents compared with the control.1-MCP treatment reduced the rate of decrease of sucrose, fructose and glucose, and maintained higher levels of sugars. When compare to control florets, the activities of sucrose phosphate synthase (SPS) and sucrose synthase-synthesis (SS-S) were enhanced in florets treated with1-MCP, whereas the activities of sucrose synthase-cleavage (SS-C), glucokinase (GK) and UDP-glucose pyrophosphorylase (UGPase) were reduced. Our results indicate that sugar may play an important role in broccoli senescence, and1-MCP may delay senescence of broccoli florets through regulating sugar metabolism and maintaining higher sugar content.
(6) The effect of a combined treatment of2.5μl/11-MCP and200mg/L6-BA on surface color, antioxidant enzymes and health-promoting compounds in broccoli florets stored at15℃was investigated. The combined treatment of1-MCP and6-BA significantly delayed senescence and maintained visual quality of broccoli florets compared with1-MCP or6-BA treatment alone during storage. The combined treatment profoundly reduced the rate of increase of L*value, and retained high level of H and chlorophyll content. The combined treatment enhanced the biosynthesis of glucosinolate and the formation of the anticarcinogen sulforaphane, which maintained the health benefit of broccoli.
(7)6-BA,1-MCP and ethanol treatments can inhibite chlorophyll degradation. When compared to control, there were no significant differences in BoCHL1, BoCHL2and BoCHL3expression in6-BA-treated broccoli florets. Ethanol treatment suppressed BoCHL2and BoCHL3expression, while1-MCP treatment induced BoCHLl, BoCHL2and BoCHL3expression. These results indicated that BoCHL1, BoCHL2and BoCHL3could be unimportant in chlorophyll degration of broccoli florets because their expression patters did not correlate with degradation of chlorophyll. The up-regulation of BoPAO was repressed by ethanol and induced by1-MCP in senescence broccoli florets. It suggest that the expression of chlorophyll degradation enzymes genes could be regulated by various postharvest treatments, leading to the suppression of florets yellowing in broccoli.
(1)研究了不同贮藏温度(273、278、283、293和303K)对青花菜在贮藏期间颜色和叶绿素含量的影响。结果表明,在283、293和303K贮藏时,青花菜-a／b值和叶绿素含量随着贮藏时间延长逐渐下降,温度越高,下降越快。而在273和278K贮藏时,-a／b值和叶绿素含量无显著变化,说明低温能有效延缓青花菜黄化和叶绿素降解。基于Arrhenius动力学方程分别建立了青花菜-a/b值、叶绿素含量与贮藏时间和温度之间的动力学模型,再根据-a/b值与叶绿素含量变化的线性关系,进一步建立了基于-a/b值的叶绿素含量预测模型。该预测模型较好的预测在273-305K温度下青花菜叶绿素含量的变化状况,为应用物理颜色参数检测方法来检测青花菜品质提供依据。
(2)研究了6-BA处理对青花菜颜色、抗氧化能力、总酚、总硫代葡萄糖苷和萝卜硫素含量的影响。结果发现200mg/L6-BA处理显著抑制了L*值和丙二醛(MDA)的增长,以及H值的下降。6-BA处理通过调控叶绿素酶和脱镁螯合酶活性来抑制叶绿素降解。与对照组相比,6-BA处理增强了DPPH自由基清除能力,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性,减弱了过氧化物酶(POD)活性。6-BA处理显著增加了总酚、总硫代葡萄糖苷和萝卜硫素含量。这些结果说明,6-BA处理能保持青花菜品质,延缓衰老和改善其营养保健价值。
(3)研究了乙醇处理对青花菜表观颜色、抗氧化酶活性和活性成分的影响。500μl/l乙醇处理显著地延长了青花菜货架寿命,抑制了L*值的上升和H值的下降。通过抑制叶绿素酶和脱镁螯合酶活性来延缓叶绿素降解。乙醇处理增强了DPPH自由基清除能力、SOD、APX和CAT活性。乙醇处理提高了总酚、总硫代葡萄糖苷和萝卜硫素含量。这些结果说明应用乙醇能延长青花菜货架期,维持感官质量和营养品质。
(4)2.5μl/11-MCP处理能有效地延长青花菜的货架寿命,抑制颜色参数H值的下降和L*值的上升,通过抑制叶绿素酶、脱镁螯合酶和POD的活性来延缓叶绿素的降解,维持其表观绿色。1-MCP处理提高了总酚含量和SOD、APX和CAT等抗氧化酶活性,使得抗氧化能力增强。同时,也延缓了总硫代葡萄糖苷和萝卜硫素含量下降,改善了青花菜的营养保健功能。
(5)离体组织培养青花菜花蕾实验结果显示,不同种类可溶性糖能抑制叶绿素的降解。研究了1-MCP处理对采后青花菜叶绿素含量、糖代谢及其相关酶活性的影响。结果表明,与对照组相比,2.5μl/l1-MCP处理组青花菜维持更高的叶绿素a和b含量。与对照组相比,1-MCP处理增强了蔗糖磷酸合酶(SPS)和蔗糖合酶合成方向(SS-S)活性,抑制了蔗糖合酶分解方向(SS-C)、葡萄糖激酶(GK)和UDP-葡萄糖焦磷酸化酶(UGPaSe)活性。1-MCP减少了蔗糖、果糖和葡萄糖含量下降,维持了较高水平糖含量。这些结果说明,糖在青花菜衰老过程中起重要作用,1-MCP处理通过调控糖代谢和维持较高水平可溶性糖来延缓青花菜衰老。
(6)研究了2.5μl/l1-MCP和200mg/L6-BA复合处理对15℃贮藏条件下青花菜表观颜色、抗氧化酶和保健成分(萝卜硫素和硫代葡萄糖苷)的影响。复合处理比单一处理更显著延缓了青花菜贮藏期间衰老的发生并维持了花蕾的品质。1-MCP复合6-BA处理保持高水平的H值和叶绿素含量,抑制了L*值的增加和POD活性,提高了CAT、APX和SOD的活性。该复合处理较单一处理更为有效地延缓硫代葡萄糖苷的分解和诱导了青花菜中萝卜硫素的形成,维持了青花菜的营养品质。
(7)6-BA、1-MCP和乙醇处理能抑制青花菜叶绿素降解,与对照组相比,6-BA处理对BocHL1、BocHL2和BocHL3的表达没有显著影响,乙醇处理抑制了BocHL2和BoCHL3的表达,而1-MCP处理却诱导了BOCHL1、BocHL2和BoCHL3的表达,这暗示着BoCHL1、BoCHL2和BoCHL3可能在叶绿素降解过程中并没有起到关键的作用。乙醇处理延缓了青花菜衰老时BoPAO正调控,1-MCP则诱导了BoPAO的表达。这些结果表明,不同采后处理通过对叶绿素降解酶基因的调控作用,从而抑制了青花菜的黄化。
Broccoli (Brassica oleracea L. ssp. italica) is known to have high nutritional value with significant contents of vitamins, glucosinolates and phenolic compounds. The inflorescences are harvested while the floral heads, branchlets, and are totally immature, with the sepals completely surrounding the flower. However, broccoli deteriorates very quickly after harvest at ambient temperature due to the loss of green color and the consequent yellowing of florets, which decreases the commercial acceptance of the broccoli florets. Therefore, it is crucial to study the regulation and mechanism of quality deterioration of broccoli florets after harvest for transportation and storage. Basing on the change of color and yellowing of broccoli florets upon various temperatures storage, the present study was designed to investigate the effect of6-benzylaminopurine (6-BA),1-methylcyclopropene (1-MCP) and ethanol on inhibiting yellowing of postharvest broccoli florets and the possible mechanisms involved. The effect of1-MCP in combination with6-BA in delaying senescence on broccoli florets was also assessed. Furthermore, the effect of1-MCP treatment on regulation sugar metabolism and involvement of enzymes in broccoli florets was investigated, which laid a theoretical basis for sugars acting as signaling molecules in regulating senescence of broccoli florets. The results were as follows:
(1) The effect of different storage temperatures (273,278,283,293and303K) on color and chlorophyll content of postharvest broccoli florets was investigated. The results showed that-a/b value and chlorophyll content of broccoli florets markedly decreased during stored at283,293and303K. The increase of storage temperature, change scope of-a/b value and chlorophyll content was also promoted. There were no significant differences in color and chlorophyll content of broccoli florets during stored at273and278K, which suggested that low temperature can delay the yellowing and chlorophyll degradation of broccoli. The kinetic models of-a/b value and chlorophyll content with respect to storage time and temperature were established based on the Arrhenius equation, respectively. Then, a prediction model for chlorophyll content based on-a/b value was developed according to the linear relationship between chlorophyll content and-a/b value. Finally, the chlorophyll content of broccoli can be accurately predicted at the storage temperature from273to305K by the prediction model, which confirms the feasibility of applying the objective color parameter measurement for assessment of broccoli quality.
(2) The effect of6-BA on color, antioxidant activity, contents of total phenols, glucosinolate and sulforaphane in broccoli florets was investigated. The results showed that200mg/L6-BA treatment markedly inhibited the increase of L value and malondialdehyde (MDA) content, and retarded the decrease of H value.6-BA treatment reduced rate of chlorophyll degradation by regulating the activities of chlorophyllase and Mg-dechelatase. When compared to control florets, the1,1-diphenyl-2-picrylhydrazyl (DPPH) radical scavenging activity and the activities of superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) were enhanced in florets treated with6-BA, whereas the activity of peroxidase (POD) was reduced. The contents of total phenols, glucosinolate and sulforaphane were increased after treated with6-BA. These results indicated that6-BA could maintain quality, delay senescence and improve the nutritional value of broccoli.
(3) The effect of cthanol treatment on visual quality, antioxidant enzymes and bioactive compounds in broccoli florets was investigated.500μ1/1ethanol treatment markedly extended shelf life, inhibited the increase of L*value and the decrease of H value. It reduced the rate of chlorophyll degradation by inhibiting the activities of chlorophyllase and Mg-dechelatase. In addition, the DPPH radical scavenging activity and the activities of SOD, APX and CAT were enhanced in florets treated with ethanol. The contents of total phenolic compounds, glucosinolates and sulforaphane were also profoundly increased after treated with ethanol. These results indicated that a application of ethanol can prolong shelf life, maintain visual quality and the nutritional value of broccoli florets.
(4)2.5μl/11-MCP treatment significantly extended shelf life, inhibited the increase of L value and the decrease of H value.1-MCP might delay chlorophyll degradation and florets yellowing by inhibiting the activities of chlorophyllase, Mg-dechelatase and POD. The enhancement in antioxidant activity by1-MCP may be due to increased phenolic content and higher activities of antioxidant enzymes including CAT, SOD and APX. Thus1-MCP has a potential application in postharvest treatment for prolonging shelf life, maintaining quality and improving the health benefit of broccoli florets.
(5) In vitro incubation of broccoli florets showed that various soluble sugars can retard the chlorophyll degradation in broccoli florets. The effect of1-methylcyclopene (1-MCP) on chlorophyll content, sugar metabolism and involvement of enzymes in broccoli florets was investigated. The results showed that1-MCP treatment maintained higher chlorophyll a and b contents compared with the control.1-MCP treatment reduced the rate of decrease of sucrose, fructose and glucose, and maintained higher levels of sugars. When compare to control florets, the activities of sucrose phosphate synthase (SPS) and sucrose synthase-synthesis (SS-S) were enhanced in florets treated with1-MCP, whereas the activities of sucrose synthase-cleavage (SS-C), glucokinase (GK) and UDP-glucose pyrophosphorylase (UGPase) were reduced. Our results indicate that sugar may play an important role in broccoli senescence, and1-MCP may delay senescence of broccoli florets through regulating sugar metabolism and maintaining higher sugar content.
(6) The effect of a combined treatment of2.5μl/11-MCP and200mg/L6-BA on surface color, antioxidant enzymes and health-promoting compounds in broccoli florets stored at15℃was investigated. The combined treatment of1-MCP and6-BA significantly delayed senescence and maintained visual quality of broccoli florets compared with1-MCP or6-BA treatment alone during storage. The combined treatment profoundly reduced the rate of increase of L*value, and retained high level of H and chlorophyll content. The combined treatment enhanced the biosynthesis of glucosinolate and the formation of the anticarcinogen sulforaphane, which maintained the health benefit of broccoli.
(7)6-BA,1-MCP and ethanol treatments can inhibite chlorophyll degradation. When compared to control, there were no significant differences in BoCHL1, BoCHL2and BoCHL3expression in6-BA-treated broccoli florets. Ethanol treatment suppressed BoCHL2and BoCHL3expression, while1-MCP treatment induced BoCHLl, BoCHL2and BoCHL3expression. These results indicated that BoCHL1, BoCHL2and BoCHL3could be unimportant in chlorophyll degration of broccoli florets because their expression patters did not correlate with degradation of chlorophyll. The up-regulation of BoPAO was repressed by ethanol and induced by1-MCP in senescence broccoli florets. It suggest that the expression of chlorophyll degradation enzymes genes could be regulated by various postharvest treatments, leading to the suppression of florets yellowing in broccoli.
引文
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