1-MCP与内源乙烯相互作用对番茄和鳄梨成熟生理的影响研究
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摘要
本文以番茄(Solanum lycopersicum L. cv. Sebring and Florida 47)和鳄梨(Persea Americana L. cv. Booth 7)果实为试材,研究了1-MCP结合外源乙烯和低压低氧处理对常温贮藏番茄和鳄梨果实成熟生理的影响,深入探讨了内源乙烯水平作为重要因素而实现调节1-MCP对跃变型果实反应效率的可能机制。主要研究结果如下:
1.首先使用气态1-MCP(516 nL L~(-1))对破色期番茄(cv. Sebring)果实处理9 h后,立即用100μL L~(-1)乙烯分别处理3 h和6 h,然后贮藏于20±1°C环境中。结果显示,1-MCP处理后的番茄果实的软化和色泽度下降受随后乙烯处理的影响较小。
与1-MCP和乙烯先后处理试验结果不同,破色期番茄(cv. Sebring)果实被100μL L~(-1)乙烯和气态1-MCP(500 nL L~(-1))同步处理6 h后,1-MCP对果实软化和色度降低的抑制能力几乎完全失去,但低浓度乙烯(10μL L~(-1))对500 nL L~(-1)1-MCP反应效率的影响甚微。
破色期番茄(cv. Florida 47)果实被100μL L~(-1)乙烯处理6 h后,分别在0 h、1 h和6 h时使用液态1-MCP(200μg L~(-1))对果实浸泡处理1 min。结果显示,1-MCP的敏感性在乙烯处理后的短时期内(0~(-1) h)被显著的降低,但在6 h后,随着内源乙烯浓度(IEC)逐渐恢复至乙烯处理前的水平,1-MCP的反应能力可以完全恢复。1-MCP恢复的反应敏感性主要表现在其对果实的软化、色度下降、呼吸跃变、乙烯释放、番茄红素合成、可滴定酸变化和总酚积累的抑制,其效率与200μg L~(-1) 1-MCP单独处理(无乙烯)相当。
2.将破色期和粉红期番茄(cv. Florida 47)果实首先在低压低氧(HH, 8 kPa, 1.7 kPa O2)条件下胁迫处理6 h,随后于常压下分别在0 h和3 h时使用液态次饱和浓度1-MCP(50μg L~(-1))对果实浸泡处理1 min。结果显示,当HH(8 kPa, 1.7 kPa O2)处理结束时,破色期和粉红期果实的内源乙烯浓度(IEC)分别较对照(无HH)降低了76%和62%,ACO活性分别较对照下降66%和60%,且果实对1-MCP的反应敏感性显著提高,提高的敏感性表现在HH+1-MCP 0 h处理较1-MCP处理更强烈地延缓了贮藏果实的硬度和色泽度下降、推迟了呼吸和乙烯释放峰和抑制了番茄红素积累。然而,在HH胁迫处理结束后3 h内,随着ACO活性和IEC升高到HH处理前水平,果实对1-MCP的反应敏感性相应地恢复到1-MCP单独处理水平。我们推测,HH调节1-MCP反应能力的提高可能归因于降低的pO2。为证明与此,将破色期和粉红期番茄(cv. Florida 47)果实首先在低压常氧(HN, 21.3 kPa, 21.3 kPa O2)条件下暴露6 h,随后于常压下在0 h时使用液态1-MCP(50μg L~(-1))对果实浸泡处理1 min。结果发现,破色期和粉红期果实的IEC和ACO活性未受HN(21.3 kPa, 21. kPa O2)处理影响。相应地,HN +1-MCP 0 h处理对果实成熟的抑制能力仅与1-MCP单独处理相当。
3.与50μg L~(-1)1-MCP单独处理果实相比,HH+1-MCP 0 h处理更为强烈地影响了多聚半乳糖醛酸酶(PG)、纤维素酶(Cx)、果胶甲脂酶(PME)、α-半乳糖苷酶(α-Gal)等细胞壁降解酶活性的变化,对PG的影响尤为明显;HH+1-MCP处理还显著地抑制了番茄果实贮藏期水溶性和CDTA溶性糖醛酸(UA)含量的增加;凝胶渗透色谱分析结果显示,1-MCP和HH+1-MCP 0 h处理延缓了番茄果实水溶性和CDTA溶性多聚糖醛酸分子的降解过程,且后者效果更为明显。
4.以100μg L~(-1)和500μg L~(-1)液态1-MCP浸泡处理强烈地抑制了跃变前(采后1 d)鳄梨果实在贮藏期的软化和色泽度下降,并推迟了呼吸和乙烯释放峰的出现。跃变中期(采后7 d)鳄梨果实完全失去了对100μg L~(-1) 1-MCP并极大减弱了对500μg L~(-1) 1-MCP的反应敏感性。然而,高剂量(2500μg L~(-1))的1-MCP处理显示出其对跃变中期果实成熟的强烈抑制。此外,低氧结合1-MCP(1000μg L~(-1))处理试验结果显示,跃变中期(采后6.5 d)鳄梨果实经空气和低氧(2 kPa O2)胁迫处理12 h后,IEC由初始值38.6μL L~(-1)分别增加和减小到54μL L~(-1)和16μL L~(-1)。较空气+1-MCP处理相比,低氧+1-MCP 0 h处理较大程度地抑制了贮藏鳄梨果实的软化、色泽度下降、呼吸速率和乙烯释放率。同时,空气+1-MCP和低氧+1-MCP处理也抑制了跃变中期果实的PG积累和水溶性多聚糖醛酸降解,且后者处理的效果更为显著,表明鳄梨果实在跃变起始后的细胞壁代谢仍依赖于乙烯作用。
上述研究结果表明,内源乙烯水平是调节1-MCP反应能力的重要因素,解释了不同跃变型果实在成熟起始后对1-MCP反应敏感性差异的原因。
Using tomato (Solanum lycopersicum L. cv. Sebring and Florida 47) and avocaodo (Persea Americana L. cv. Booth 7) fruit as a model system, this study tested the idea that internal ethylene levels can modulate the efficacy of 1-MCP at suppressing ripening in climacteric fruits. The main results were exhibited as follows:
1. Breaker tomato (cv. Sebring) fruit were treated with gaseous 1-MCP (SmartFreshSM Quality System) under conditions (516 nL L~(-1)) affording maximum inhibition of ripening, followed by subsequent exposure to 100μL~(-1) ethylene for 3 or 6 h. Fruit softening and hue angle decline in 1-MCP-treated fruit were minimally affected in response to ethylene。In contrast to sequential 1-MCP and ethylene treatments, simultaneous treatment of breaker‘Sebring’tomato fruit with 100μL L~(-1) ethylene and gaseous 500 nL L~(-1) 1-MCP completely negated the capacity of 1-MCP to inhibit fruit softening and hue angle decline. When breaker fruit were treated with 100μL L ~(-1) ethylene for 6 h followed by exposure to aqueous 1-MCP (200μg L~(-1)), sensitivity to 1-MCP was significantly reduced in the short-term (0–1 h) and recovered within 6 h to patterns characteristic of fruit receiving 200μg L~(-1) aqueous 1-MCP without prior exposure to ethylene. Re-sensitization was reflected in patterns of softening, climacteric ethylene and respiratory responses, hue angle decline, lycopene content, titratable acidity and total phenolic content changes. The time required for re-sensitization to 1-MCP paralleled the time required for return of internal ethylene levels to concentrations present prior to ethylene treatment.
2. Breaker and pink‘Florida 47’tomato fruit were subjected to hypobaria hypoxic (HH) (8 kPa, 1.7 kPa O2) for 6 h, followed by treatment at ambient pressure with aqueous 1-MCP at a sub-saturating dose (50μg L~(-1)) and exposure duration (1 min). Immediately following HH, breaker and pink fruit had a 76% and 62% lower IEC, 66% and 60% lower ACO activities, respectively, as well as enhanced sensitivity to 1-MCP compared with fruit not receiving HH. Increased sensitivity to 1-MCP was evident in further suppression of fruit firmness and hue angle declines, and delayed peak ethylene production. Within 3 h of removal of fruit from HH, IEC and ACO activity had returned to pre-HH values and responses to 1-MCP were comparable with those of fruit receiving 1-MCP without prior HH. Decreased IEC and increased sensitivity to 1-MCP in response to HH were also observed for fruit at more advanced ripening (pink stage). We addressed whether increased sensitivity to 1-MCP following exposure to HH was a response to reduced pO2. For this objective, breaker and pink fruit were exposed to hypobaria normoxic (HN) (21.3 kPa, 21.3 kPaO2) for 6 h prior to treatment with 50μg L?1 aqueous 1-MCP. The results showed that IEC, ACO activity and responsiveness to 1-MCP were unaffected in fruit following exposure to HN, indicating that enhanced sensitivity to 1-MCP following HH is a result of low pO2-mediated reductions in IEC.
3. The immediate treatment with 1- MCP after HH (8 kPa, 1.7 kPa O2) also more evidently affected the activity trends of cell wall enzymes including polygalacturonase (PG), cellulase (Cx) pectinmethylesterase (PME),α-galactosidase (α-Gal) activity, typically on PG activity. Consistent with trend of PG activity, water/CDTA-soluble uronic acid (UA) content were more strongly suppressed by treatment of 50μg L~(-1) 1-MCP following HH, compared with 1-MCP treatment without prior to HH; Polyuronides in fruit treated with 1-MCP following HH exhibited slower molecular mass downshifts compared with the control and 1-MCP- treated fruit. In addition, all results in this study showed that there was no significant difference between control and HH treatment.
4. The role of IEC in modulating 1-MCP responsiveness was further tested using avocado, a fruit that accumulates markedly higher IEC compared with tomato fruit. Preclimacteric (1 d after harvest)‘Booth 7’avocado fruit were treated for 1 min with aqueous 1-MCP at 100 and 500μg L~(-1). Both concentrations strongly suppressed softening, and delayed climacteric ethylene production and respiration maxima. Mid-climacteric fruit (7 d after harvest) showed complete loss of or diminished sensitivity to 1-MCP at 100 and 500μg L~(-1), respectively. Application of higher dose of aqueous 1-MCP (2500μg L~(-1)) revealed that softening, respiration and ethylene production of mid-climacteric avocado remained highly sensitive to inhibition of ethylene perception. In experiments testing the effects of reducing IEC on 1-MCP responses, mid-climacteric (d 6.5) avocado were exposed to hypoxia (2 kPa O2) for 12 h, followed by exposure to aqueous 1-MCP at 1000μg L~(-1). IEC prior to hypoxia averaged about 38.6μL L~(-1). IEC following exposure to air or hypoxia for 12 h averaged around 54 and 16μL L~(-1), respectively. Fruit treated with 1000μg L~(-1)1-MCP following hypoxia showed greater suppression of fruit softening, and further delays in peak ethylene production and respiration compared with fruit treated with 1-MCP alone. 1-MCP and low O2+1-MCP treatments applied to mid-climacteric fruit also delayed PG accumulation and depolymerization of polyuronides, indicating that cell wall metabolism in avocado fruit remains ethylene sensitive through advanced ripening. Possible mechanisms by which 1-MCP sensitivity is modulated by ethylene are discussed.
In summary, we propose that internal ethylene levels may contribute to the divergent sensitivities of some climacteric fruits to 1-MCP applied after initiation of ripening.
1.首先使用气态1-MCP(516 nL L~(-1))对破色期番茄(cv. Sebring)果实处理9 h后,立即用100μL L~(-1)乙烯分别处理3 h和6 h,然后贮藏于20±1°C环境中。结果显示,1-MCP处理后的番茄果实的软化和色泽度下降受随后乙烯处理的影响较小。
与1-MCP和乙烯先后处理试验结果不同,破色期番茄(cv. Sebring)果实被100μL L~(-1)乙烯和气态1-MCP(500 nL L~(-1))同步处理6 h后,1-MCP对果实软化和色度降低的抑制能力几乎完全失去,但低浓度乙烯(10μL L~(-1))对500 nL L~(-1)1-MCP反应效率的影响甚微。
破色期番茄(cv. Florida 47)果实被100μL L~(-1)乙烯处理6 h后,分别在0 h、1 h和6 h时使用液态1-MCP(200μg L~(-1))对果实浸泡处理1 min。结果显示,1-MCP的敏感性在乙烯处理后的短时期内(0~(-1) h)被显著的降低,但在6 h后,随着内源乙烯浓度(IEC)逐渐恢复至乙烯处理前的水平,1-MCP的反应能力可以完全恢复。1-MCP恢复的反应敏感性主要表现在其对果实的软化、色度下降、呼吸跃变、乙烯释放、番茄红素合成、可滴定酸变化和总酚积累的抑制,其效率与200μg L~(-1) 1-MCP单独处理(无乙烯)相当。
2.将破色期和粉红期番茄(cv. Florida 47)果实首先在低压低氧(HH, 8 kPa, 1.7 kPa O2)条件下胁迫处理6 h,随后于常压下分别在0 h和3 h时使用液态次饱和浓度1-MCP(50μg L~(-1))对果实浸泡处理1 min。结果显示,当HH(8 kPa, 1.7 kPa O2)处理结束时,破色期和粉红期果实的内源乙烯浓度(IEC)分别较对照(无HH)降低了76%和62%,ACO活性分别较对照下降66%和60%,且果实对1-MCP的反应敏感性显著提高,提高的敏感性表现在HH+1-MCP 0 h处理较1-MCP处理更强烈地延缓了贮藏果实的硬度和色泽度下降、推迟了呼吸和乙烯释放峰和抑制了番茄红素积累。然而,在HH胁迫处理结束后3 h内,随着ACO活性和IEC升高到HH处理前水平,果实对1-MCP的反应敏感性相应地恢复到1-MCP单独处理水平。我们推测,HH调节1-MCP反应能力的提高可能归因于降低的pO2。为证明与此,将破色期和粉红期番茄(cv. Florida 47)果实首先在低压常氧(HN, 21.3 kPa, 21.3 kPa O2)条件下暴露6 h,随后于常压下在0 h时使用液态1-MCP(50μg L~(-1))对果实浸泡处理1 min。结果发现,破色期和粉红期果实的IEC和ACO活性未受HN(21.3 kPa, 21. kPa O2)处理影响。相应地,HN +1-MCP 0 h处理对果实成熟的抑制能力仅与1-MCP单独处理相当。
3.与50μg L~(-1)1-MCP单独处理果实相比,HH+1-MCP 0 h处理更为强烈地影响了多聚半乳糖醛酸酶(PG)、纤维素酶(Cx)、果胶甲脂酶(PME)、α-半乳糖苷酶(α-Gal)等细胞壁降解酶活性的变化,对PG的影响尤为明显;HH+1-MCP处理还显著地抑制了番茄果实贮藏期水溶性和CDTA溶性糖醛酸(UA)含量的增加;凝胶渗透色谱分析结果显示,1-MCP和HH+1-MCP 0 h处理延缓了番茄果实水溶性和CDTA溶性多聚糖醛酸分子的降解过程,且后者效果更为明显。
4.以100μg L~(-1)和500μg L~(-1)液态1-MCP浸泡处理强烈地抑制了跃变前(采后1 d)鳄梨果实在贮藏期的软化和色泽度下降,并推迟了呼吸和乙烯释放峰的出现。跃变中期(采后7 d)鳄梨果实完全失去了对100μg L~(-1) 1-MCP并极大减弱了对500μg L~(-1) 1-MCP的反应敏感性。然而,高剂量(2500μg L~(-1))的1-MCP处理显示出其对跃变中期果实成熟的强烈抑制。此外,低氧结合1-MCP(1000μg L~(-1))处理试验结果显示,跃变中期(采后6.5 d)鳄梨果实经空气和低氧(2 kPa O2)胁迫处理12 h后,IEC由初始值38.6μL L~(-1)分别增加和减小到54μL L~(-1)和16μL L~(-1)。较空气+1-MCP处理相比,低氧+1-MCP 0 h处理较大程度地抑制了贮藏鳄梨果实的软化、色泽度下降、呼吸速率和乙烯释放率。同时,空气+1-MCP和低氧+1-MCP处理也抑制了跃变中期果实的PG积累和水溶性多聚糖醛酸降解,且后者处理的效果更为显著,表明鳄梨果实在跃变起始后的细胞壁代谢仍依赖于乙烯作用。
上述研究结果表明,内源乙烯水平是调节1-MCP反应能力的重要因素,解释了不同跃变型果实在成熟起始后对1-MCP反应敏感性差异的原因。
Using tomato (Solanum lycopersicum L. cv. Sebring and Florida 47) and avocaodo (Persea Americana L. cv. Booth 7) fruit as a model system, this study tested the idea that internal ethylene levels can modulate the efficacy of 1-MCP at suppressing ripening in climacteric fruits. The main results were exhibited as follows:
1. Breaker tomato (cv. Sebring) fruit were treated with gaseous 1-MCP (SmartFreshSM Quality System) under conditions (516 nL L~(-1)) affording maximum inhibition of ripening, followed by subsequent exposure to 100μL~(-1) ethylene for 3 or 6 h. Fruit softening and hue angle decline in 1-MCP-treated fruit were minimally affected in response to ethylene。In contrast to sequential 1-MCP and ethylene treatments, simultaneous treatment of breaker‘Sebring’tomato fruit with 100μL L~(-1) ethylene and gaseous 500 nL L~(-1) 1-MCP completely negated the capacity of 1-MCP to inhibit fruit softening and hue angle decline. When breaker fruit were treated with 100μL L ~(-1) ethylene for 6 h followed by exposure to aqueous 1-MCP (200μg L~(-1)), sensitivity to 1-MCP was significantly reduced in the short-term (0–1 h) and recovered within 6 h to patterns characteristic of fruit receiving 200μg L~(-1) aqueous 1-MCP without prior exposure to ethylene. Re-sensitization was reflected in patterns of softening, climacteric ethylene and respiratory responses, hue angle decline, lycopene content, titratable acidity and total phenolic content changes. The time required for re-sensitization to 1-MCP paralleled the time required for return of internal ethylene levels to concentrations present prior to ethylene treatment.
2. Breaker and pink‘Florida 47’tomato fruit were subjected to hypobaria hypoxic (HH) (8 kPa, 1.7 kPa O2) for 6 h, followed by treatment at ambient pressure with aqueous 1-MCP at a sub-saturating dose (50μg L~(-1)) and exposure duration (1 min). Immediately following HH, breaker and pink fruit had a 76% and 62% lower IEC, 66% and 60% lower ACO activities, respectively, as well as enhanced sensitivity to 1-MCP compared with fruit not receiving HH. Increased sensitivity to 1-MCP was evident in further suppression of fruit firmness and hue angle declines, and delayed peak ethylene production. Within 3 h of removal of fruit from HH, IEC and ACO activity had returned to pre-HH values and responses to 1-MCP were comparable with those of fruit receiving 1-MCP without prior HH. Decreased IEC and increased sensitivity to 1-MCP in response to HH were also observed for fruit at more advanced ripening (pink stage). We addressed whether increased sensitivity to 1-MCP following exposure to HH was a response to reduced pO2. For this objective, breaker and pink fruit were exposed to hypobaria normoxic (HN) (21.3 kPa, 21.3 kPaO2) for 6 h prior to treatment with 50μg L?1 aqueous 1-MCP. The results showed that IEC, ACO activity and responsiveness to 1-MCP were unaffected in fruit following exposure to HN, indicating that enhanced sensitivity to 1-MCP following HH is a result of low pO2-mediated reductions in IEC.
3. The immediate treatment with 1- MCP after HH (8 kPa, 1.7 kPa O2) also more evidently affected the activity trends of cell wall enzymes including polygalacturonase (PG), cellulase (Cx) pectinmethylesterase (PME),α-galactosidase (α-Gal) activity, typically on PG activity. Consistent with trend of PG activity, water/CDTA-soluble uronic acid (UA) content were more strongly suppressed by treatment of 50μg L~(-1) 1-MCP following HH, compared with 1-MCP treatment without prior to HH; Polyuronides in fruit treated with 1-MCP following HH exhibited slower molecular mass downshifts compared with the control and 1-MCP- treated fruit. In addition, all results in this study showed that there was no significant difference between control and HH treatment.
4. The role of IEC in modulating 1-MCP responsiveness was further tested using avocado, a fruit that accumulates markedly higher IEC compared with tomato fruit. Preclimacteric (1 d after harvest)‘Booth 7’avocado fruit were treated for 1 min with aqueous 1-MCP at 100 and 500μg L~(-1). Both concentrations strongly suppressed softening, and delayed climacteric ethylene production and respiration maxima. Mid-climacteric fruit (7 d after harvest) showed complete loss of or diminished sensitivity to 1-MCP at 100 and 500μg L~(-1), respectively. Application of higher dose of aqueous 1-MCP (2500μg L~(-1)) revealed that softening, respiration and ethylene production of mid-climacteric avocado remained highly sensitive to inhibition of ethylene perception. In experiments testing the effects of reducing IEC on 1-MCP responses, mid-climacteric (d 6.5) avocado were exposed to hypoxia (2 kPa O2) for 12 h, followed by exposure to aqueous 1-MCP at 1000μg L~(-1). IEC prior to hypoxia averaged about 38.6μL L~(-1). IEC following exposure to air or hypoxia for 12 h averaged around 54 and 16μL L~(-1), respectively. Fruit treated with 1000μg L~(-1)1-MCP following hypoxia showed greater suppression of fruit softening, and further delays in peak ethylene production and respiration compared with fruit treated with 1-MCP alone. 1-MCP and low O2+1-MCP treatments applied to mid-climacteric fruit also delayed PG accumulation and depolymerization of polyuronides, indicating that cell wall metabolism in avocado fruit remains ethylene sensitive through advanced ripening. Possible mechanisms by which 1-MCP sensitivity is modulated by ethylene are discussed.
In summary, we propose that internal ethylene levels may contribute to the divergent sensitivities of some climacteric fruits to 1-MCP applied after initiation of ripening.
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