外源油菜素内酯对改善骏枣果实品质及减轻裂果的效果

杨 成,王振磊,张川疆,刘 伟,吴翠云,林敏娟*

(塔里木大学园艺与林学学院·南疆特色果树高效优质栽培与深加工技术国家地方联合工程实验室·塔里木盆地生物资源保护利用省部共建国家重点实验室培育基地·塔里木绿洲农业教育部重点实验室,新疆阿拉尔 843300)

摘 要:【目的】探究不同浓度油菜素内酯(BR)处理对骏枣果实裂果及相关生理特性的影响。【方法】以10年生骏枣为试材,在果实膨大期喷施不同质量浓度BR(1、2、3 mg·L-1),研究BR对骏枣裂果率、果实品质、细胞壁代谢酶活性及内源激素含量的影响。【结果】不同浓度BR处理显著降低了裂果率,各处理裂果关键期都处于白熟期,全红期为裂果高峰期。不同浓度BR处理在全红期较对照果实裂果率分别降低了6.38%、4.63%、4.48%,不同浓度BR处理提高了果实的营养品质,提高了果实硬度、咀嚼性和胶黏性;全红期的枣果皮CAT 活性分别比对照提高了18.54%、39.07%、98.01%,SOD 活性分别提高了55.32%、67.25%、71.18%,POD 活性分别降低了24.66%、1.15%、32.06%,纤维素酶活性及果胶酶相较于对照降低,其中,果胶酶活性比对照分别降低了6.92%、7.75%、13.91%;BR处理对GA3和IAA含量在各时期都具有提高效果,全红期GA3含量较对照分别提高了8.30%、24.70%和14.07%。IAA含量分别提高了312.12%、198.48%、457.58%,对ABA含量具有降低作用。【结论】喷施BR能有效降低骏枣裂果率,改善果实品质,提高果皮抗氧化酶活性及GA3含量和IAA含量,降低纤维素酶、果胶酶活性及ABA含量,减少裂果发生。其中,1 mg·L-1的油菜素内酯对骏枣防裂效果最佳。

关键词:骏枣;油菜素内酯;裂果;抗氧化酶活性

枣(Ziziphus jujuba Mill.)为鼠李科枣属植物,是新疆第一大林果产业,现已形成环塔里木盆地林果产区,栽培面积达33万hm2[1]。裂果在石榴[2]、番茄[3]、葡萄[4]等上都有发生。长期以来枣裂果的问题始终困扰着红枣产业的发展,随着果实成熟度的增加及雨水等因素的影响,裂果率达到50%以上,甚至于绝收,严重降低了果实品质及商品价值,制约着枣产业发展[5-6]

果实遇到逆境胁迫时会产生一些活性氧物质,当这些活性氧物质高于抗氧化酶的清除能力时,就会加速植物的膜脂过氧化反应,破坏细胞膜系统,对果实细胞造成伤害,从而诱发裂果,抗裂果实内的抗氧化酶活性也会高于易裂果实或已裂果实[7-9]。纤维素酶和果胶酶作为分解细胞壁物质的酶类,此类酶活性的高低与裂果有密切的关系,在易裂果荔枝品种中的活性显著高于抗裂果品种[10-11]。质地是果实品质的重要组成部分,是影响果实裂果的重要因素之一[12]。前人在研究过程中发现,硬度是影响裂果的主要因素,对不同樱桃品种研究也说明了硬度越高的甜樱桃品种更容易开裂[13-14]

油菜素内酯(BR)是一种广泛存在于植物体内的激素,可以增加果皮和细胞壁的强度,提高果实抗裂能力和果实品质[15-17]。研究表明,BR 能提高果实的营养品质,增加果实硬度,提高番茄果皮抗氧化酶活性,降低细胞壁水解酶活性,从而有效降低番茄裂果率[18]。在荔枝的防裂研究中发现,1 mg·L-1 BR显著降低了荔枝裂果率,提高了果实的营养品质[8]。目前,有关油菜素内酯对枣裂果影响的研究鲜有报道,因此笔者在本研究中以骏枣为试材,研究油菜素内酯对骏枣不同发育时期果实裂果率、果实品质、相关酶活性和内源激素含量的影响,通过相关性分析及主成分分析,探讨果实裂果与相关性状的关系,以期为新疆枣裂果防控提供参考。

1 材料和方法

1.1 材料

试验所在地阿拉尔市2024年降水量为83.8 mm,主要集中在8 月,降雨量为63.4 mm。试验于2024年6—11月在新疆生产建设兵团第一师十团三连进行,供试材料为树势良好、栽培管理条件一致的10年生骏枣树(图1)。

图1 白熟期裂果骏枣
Fig.1 Junzao with cracked fruit at the white ripening period

1.2 试验设计

采用单因素完全随机试验设计,设置3 个处理及1 个对照,每个处理9 株树,分3 次重复。在2024年6月对枣树进行挂牌标记,从果实膨大期(2024年7 月17 日)开始喷施不同质量浓度的BR(分别为1、2、3 mg·L-1)及清水对照(CK)。采用叶面喷施,每隔7 d 喷施1 次,共计3 次,以枣果及叶面向下滴水为度,均在20:00开始喷施。

于果实绿熟期、白熟期、转色期、半红期、全红期采摘大小均一、无裂果、无机械损伤、无病虫害的果实各60个,对其中30个果实分离果皮、果肉,果皮液氮冷冻后置于超低温冰箱保存,用于测定内源激素含量及细胞壁代谢酶活性,10个果实用于测定果实质地,20个果实用于测定果实品质。

1.3 试验方法

自然裂果率调查:于果实不同生长发育期对自然裂果率进行调查,对标记的未取样的枝条调查总果数、裂果数。

累积裂果率/%=(裂果个数/调查总果实数)×100。

各时期裂果率/%=每时期裂果率-前一时期裂果率。

采用PAL-1 手持折光仪测定可溶性固形物含量,可溶性糖含量测定采用蒽酮硫酸法,可滴定酸含量测定采用酸碱滴定法。

果实质地测定:使用美国FTC 公司TMS-PRO质构仪。测定果实硬度、粘附性、弹性、胶黏性、咀嚼性等,质构仪设置参数参考杨植等[13]的方法。

枣果皮相关酶活性测定:过氧化氢酶(CAT)活性测定采用紫外吸收法,过氧化物酶(POD)活性测定采用愈创木酚法[11,19],超氧化物歧化酶(SOD)活性测定采用苏州格瑞斯生物科技有限公司WST-8法活性测定试剂盒。纤维素酶及果胶酶活性测定采用DNS终止法[20]

枣果皮内源激素含量的测定:GA3、IAA及ABA含量的测定参考杨途熙等[21]的方法。

1.4 数据分析

使用Microsoft Excel 2016 进行数据收集整理,采用Origin 2021进行图表绘制、相关性分析及主成分分析,采用SPSS 27.0进行方差分析及差异显著性分析。

2 结果与分析

2.1 施用不同浓度油菜素内酯对骏枣自然裂果率的影响

由图2可知,骏枣果实裂果率随果实成熟而升高,于果实绿熟期发生裂果,在全红期达到最高。白熟期、转色期、半红期、全红期骏枣累积裂果率都极显著低于对照,各时期裂果率中,处理及对照的裂果率在白熟期最高,分别为7.05%、7.43%、7.64%和11.79%,因此,在降雨量增高的情况下,施用BR的骏枣果实较对照更加抗裂,各BR 处理间抗裂效果接近。随着成熟度增加,裂果率增速降低,因此认为骏枣的裂果关键期为白熟期。在果实全红期时,BR处理累积裂果率较对照分别极显著降低了6.38%、4.63%、4.48%,其中,以1 mg·L-1 BR处理抗裂效果最佳。

图2 油菜素内酯对骏枣裂果率的影响
Fig.2 Effects of brassinolide on fruit cracking rate of Junzao

2.2 施用不同浓度油菜素内酯对骏枣果实品质的影响

由图3-A 可知,可溶性固形物含量随果实成熟度增加而上升,喷施BR 对骏枣可溶性固形物含量有提升作用,其中绿熟期、转色期(2 mg·L-1 BR处理除外)可溶性固形物含量均显著高于对照,在全红期时,3 mg·L-1 BR处理的果实可溶性固形物含量极显著高于对照,提高9.5%。

图3 油菜素内酯对不同时期骏枣果实品质的影响
Fig.3 Effects of brassinolide on fruit quality of Junzao in different periods

由图3-B 可知,可溶性糖含量随果实成熟度增加而上升,喷施BR可以提高果实可溶性糖含量,在绿熟期时,2 mg·L-1 BR处理较对照显著提高3.42%,在全红期各处理可溶性糖含量与对照无显著差异。

由图3-C可知,与对照相比,绿熟期、半红期各处理可滴定酸含量显著降低,其余生长发育时期无显著差异,除转色期和全红期1 mg·L-1 BR处理外,可滴定酸含量均为处理组低于对照。

2.3 施用不同浓度油菜素内酯对骏枣果实质地的影响

由表1 可知,在骏枣生长发育期内,半红期枣果实硬度最高,其中3 mg·L-1 BR 处理较对照显著提高了16.76%,达371.81 N,绿熟期果实硬度最低,且1 mg·L-1 BR 处理显著低于对照,在白熟期、转色期、半红期时,各处理硬度高于对照,在全红期裂果率最低的1 mg·L-1 BR 处理硬度较对照提高了9.56%。生长发育时期内果实粘附性差异较小,于果实半红期整体表现最小。果实弹性随生长发育变化整体上呈升高的趋势,在白熟期时弹性较对照分别增加了4.39%、11.11%、8.48%,在全红期3 mg·L-1 BR处理相较于对照降低了0.92%,各处理间无显著变化。各处理果实胶黏性自白熟期开始高于对照,白熟期3 mg·L-1 BR 处理较对照显著提高了24.75%,至果实全红期时,1 mg·L-1 BR 处理显著高于对照,其余处理较上一时期降低,与对照无显著差异。白熟期3 mg·L-1 BR 处理咀嚼性显著高于对照,转色期2 mg·L-1 BR 处理咀嚼性显著高于对照,半红期及全红期表现一致,各处理与对照均无显著差异。

表1 油菜素内酯对不同时期骏枣果实质地的影响
Table 1 Effects of brassinolide on fruit texture of Junzao in different periods

注:同列不同小写字母表示差异显著(p<0.05),不同大写字母表示差异极显著(p<0.01)。
Note:Different small letters in the same column indicate significant difference(p<0.05)and different capital letters indicate extremely significant difference(p<0.01).

采样时期Sampling period绿熟期Green ripening period白熟期White ripening period转色期Color change period半红期Semi-red period全红期All-red period咀嚼性Chewiness/mJ 353.68±104.37 bA 372.67±130.83 bA 528.40±115.52 aA 541.94±193.61 aA 490.18±138.53 abA 515.42±107.29 abA 596.79±141.07 aA 435.33±70.17 bA 438.84±83.96 bB 615.50±103.34 aA 465.11±89.65 bB 473.34±132.09 bB 606.18±99.57 aA 644.12±154.19 aA 644.77±194.50 aA 502.57±170.65 aA 771.70±192.22 aA 800.41±136.76 aA 765.53±176.33 aA 686.71±133.45 aA处理Treatment 1 mg·L-1 2 mg·L-1 3 mg·L-1对照CK 1 mg·L-1 2 mg·L-1 3 mg·L-1对照CK 1 mg·L-1 2 mg·L-1 3 mg·L-1对照CK 1 mg·L-1 2 mg·L-1 3 mg·L-1对照CK 1 mg·L-1 2 mg·L-1 3 mg·L-1对照CK硬度Hardness/N 233.35±37.51 cB 249.65±40.94 bcAB 300.90±38.08 aA 280.19±46.85 abAB 282.89±67.17 abA 293.83±38.20 abA 330.50±50.28 aA 271.74±44.19 bA 289.04±35.86 bA 340.18±33.75 aA 302.26±49.35 abA 282.00±48.64 bA 362.39±31.40 abA 358.73±48.95 abA 371.81±59.73 aA 318.45±43.73 bA 346.19±80.12 aA 340.23±35.11 aA 315.84±74.97 aA 315.98±31.58 aA粘附性Adhesiveness/(N·mm-1)0.06±0.02 bA 0.06±0.02 bA 0.10±0.07 aA 0.07±0.03 bA 0.07±0.03 bA 0.07±0.03 bA 0.11±0.04 aA 0.08±0.04 abA 0.05±0.03 bA 0.09±0.04 aA 0.06±0.04 abA 0.08±0.04 abA 0.03±0.01 aA 0.04±0.02 aA 0.04±0.02 aA 0.05±0.03 aA 0.08±0.05 aA 0.06±0.03 aA 0.08±0.05 aA 0.07±0.04 aA弹性Springness/mm 3.02±0.47 bB 3.04±0.49 bB 3.52±0.38 aAB 3.88±0.54 aA 3.57±0.30 abA 3.80±0.28 aA 3.71±0.35 abA 3.42±0.22 bA 3.36±0.27 bB 3.89±0.34 aA 3.33±0.21 bB 3.70±0.40 aAB 4.06±0.27 aA 4.30±0.47 aA 4.03±0.43 aA 3.91±0.49 aA 5.17±0.42 aA 5.52±0.42 aA 5.41±0.41 aA 5.46±0.53 aA胶黏性Gumminess/N 114.97±18.40 bA 118.74±24.47 bA 148.47±19.77 aA 136.19±32.47 abA 136.74±31.99 abA 134.49±21.74 abA 159.03±24.86 aA 127.48±19.97 bA 130.01±17.78 bB 157.41±13.48 aA 139.27±21.84 abAB 126.06±22.17 bB 148.84±17.37 abA 149.44±27.80 abA 157.08±33.54 aA 125.88±28.70 bA 150.12±38.91 aA 144.42±16.58 abA 142.62±34.78 abA 125.55±19.93 bA

2.4 施用不同浓度油菜素内酯对骏枣果皮细胞壁相关酶活性的影响

由图4-A可知,BR处理的骏枣果皮中CAT活性整体上随着果实成熟度的增加而呈现下降的趋势,在绿熟期、白熟期,各处理均显著高于对照,BR处理在不同发育时期相较于对照都处于高水平,说明喷施BR能提高骏枣果皮CAT活性。

图4 油菜素内酯对不同时期骏枣果皮细胞壁代谢酶活性的影响
Fig.4 Effects of brassinolide on cell wall metabolic enzyme activity of Junzao peel in different periods

由图4-B 可知,各处理组POD 活性在生长发育期呈现先上升后下降的趋势,对照则呈现上升趋势,在裂果关键期,BR 处理的骏枣果皮中POD 活性都极显著高于对照,随着成熟度的增加,各处理POD活性在全红期时低于对照,其中1、3 mg·L-1 BR处理显著低于对照。

由图4-C 可知,不同BR 处理SOD 活性随果实成熟度增加呈不同的变化趋势,除转色期和绿熟期2 mg·L-1 BR处理外,喷施BR的骏枣果皮中SOD活性在各时期均显著高于对照。在裂果关键期(白熟期)时,各处理的SOD 活性较对照分别极显著提高了43.81%、41.81%、35.08%,全红期各处理的SOD 活性较对照分别极显著提高了55.32%、67.25%、71.18%,说明喷施BR能提高骏枣果皮SOD活性。

由图4-D、E 可知,纤维素酶和果胶酶活性随果实成熟度的增加整体上呈先上升后下降的趋势,除果实全红期外,BR 处理的纤维素酶活性均低于对照,果胶酶活性在各时期均表现为处理低于对照,且在转色期和全红期,各处理的果胶酶活性均显著低于对照。说明BR 可以通过调节细胞壁代谢酶活性,从而降低果实裂果率。

2.5 施用不同浓度油菜素内酯对骏枣果皮内源激素含量的影响

由图5-A 可知,骏枣果皮GA3含量随着成熟度的提高整体上呈先上升后下降的趋势,在白熟期各处理达到峰值,其中喷施1 mg·L-1 BR时GA3含量最高,达32.34 ng·g-1,较对照极显著提高了22.18%,全红期各处理GA3含量最低,各处理在生长发育期内GA3含量均显著高于对照。

图5 油菜素内酯对不同时期骏枣果皮内源激素含量的影响
Fig.5 Effects of brassinolide on endogenous hormones content of Junzao peel in different periods

由图5-B可知,随着果实成熟期的增加,IAA含量整体呈下降趋势,在白熟期时,对照果皮中IAA含量最高,为25.67 ng·g-1,在果实半红期(2 mg·L-1 BR处理除外)及全红期,BR处理中IAA含量显著高于对照,全红期比对照分别极显著提高了312.12%、198.48%、457.58%。

由图5-C 可知,ABA 含量随着果实成熟度的增加呈现上升-下降-上升的趋势,在果实转色期时ABA 含量降低,且1 mg·L-1 BR 处理的骏枣果皮中ABA含量极显著高于对照,其他时期各处理组ABA含量显著低于对照,各组在全红期达到峰值,ABA含量(w,后同)分别为2 849.21、2 639.14、2 822.04及3 404.22 ng·g-1,说明喷施BR 对骏枣果皮脱落酸含量有降低的效果。

在果实全红期时,BR 处理及对照的(GA3+IAA)/ABA 比值分别为49、57、55、33,各处理(GA3+IAA)/ABA值均高于对照。

2.6 白熟期骏枣裂果性状相关性分析

通过对不同时期各处理裂果率与对照比较,分析得出白熟期为裂果关键期,因此,对白熟期果实裂果率、果实品质、细胞壁相关酶活性及内源激素含量等性状进行相关性分析。由图6 可知,骏枣裂果率与可滴定酸含量呈极显著正相关,与ABA含量呈显著正相关,与硬度、粘附性、CAT 活性及GA3含量呈极显著负相关,与SOD 活性呈显著负相关,表明可滴定酸含量、脱落酸含量越高,骏枣果实裂果率越高,硬度、粘附性、SOD活性、CAT活性及GA3含量越高,裂果率越低。

图6 白熟期骏枣17 个果实性状相关性分析
Fig.6 Correlation analysis of 17 fruit traits of Junzao in white ripe stage

2.7 白熟期骏枣裂果性状主成分分析

由图7 可知,对裂果关键期骏枣性状进行主成分分析,第一主成分占总变异系数的34.0%,第二主成分占总变异系数的24.8%,其中,第一主成分主要为SOD 活性、裂果率、GA3含量,为裂果因子,第二主成分主要是硬度、咀嚼性、粘附性,为硬度因子。

图7 白熟期骏枣17 个果实性状主成分分析及各元素对PC1 和PC2 的负荷
Fig.7 Principal component analysis of 17 fruit traits and the load of each element on PC1 and PC2 in Junzao at white maturity stag

3 讨 论

油菜素内酯(BR)是一种具有广谱、高效、无毒的激素,具有提高果实品质、显著影响细胞壁酶活性、提高果实内源激素含量、提高果皮韧性等优点[22]

3.1 油菜素内酯对骏枣裂果率的影响

自白熟期开始,随着降雨量增多、降雨时间增长,枣裂果情况越严重[23-24],笔者在本试验中发现白熟期为骏枣裂果关键期,裂果率增高的研究结果与之一致。喷施油菜素内酯的骏枣果实在白熟期的裂果率增速低于清水对照,表明BR 能提高骏枣的抗裂能力,在全红期累积裂果率中,各处理裂果率比对照分别降低了6.38%、4.63%及4.48%,与杨俊强等[7]在枣、Ghosh等[8]在荔枝上的研究结果一致。骏枣裂果率与可滴定酸及脱落酸含量呈显著正相关,与CAT 活性、SOD 活性、GA3含量、硬度及粘附性呈显著负相关,说明影响枣果实裂果率有许多因素,且裂果与细胞壁代谢酶活性、果实品质及内源激素含量具有较高的关联性。

3.2 油菜素内酯对骏枣果实品质的影响

施用BR 能提高骏枣可溶性糖、可溶性固形物含量及糖酸比,其中3 mg·L-1 BR处理对可溶性固形物含量提升效果最佳,其余处理对果实可溶性固形物、可溶性糖含量有提高效果,差异不显著,说明BR对骏枣果实可溶性固形物、可溶性糖含量有提高作用,但针对BR 提高骏枣果实品质的最适浓度还需进一步研究。前人也证实了对葡萄[25]、桃[26]等水果施用适宜浓度的BR 可以提高果实品质。目前,针对施用BR 提高枣果实品质的研究应用前景很广阔,笔者认为施用BR 对枣果实抗裂能力及果实品质都具有提高作用,可以在未来开展更深入的研究。

骏枣果实生长期内果实硬度整体呈先上升后下降的趋势,半红期时果实硬度最高;果实弹性、咀嚼性整体呈上升趋势,与院钦等[27]在灵武长枣上的研究结果相似。油菜素内酯处理在白熟期后对骏枣果实硬度、胶黏性具有提高的作用。对果实不同性状进行相关性分析表明,硬度、粘附性高的果实抗裂能力更强。通过主成分分析得出,硬度、咀嚼性及粘附性为第二主成分,在整个性状中占比高,是影响裂果的主要因素之一,这一结论与杨植等[13]对枣裂果与硬度关系的研究一致。

3.3 油菜素内酯对骏枣果皮细胞壁酶活性的影响

果实裂果是一种处于逆境的过程,果皮内部的活性氧及超氧阴离子产出增加,CAT、SOD作为普遍存在的抗氧化酶,可以清除活性氧及超氧阴离子,保护细胞膜组织,达到减少裂果的目的。研究表明,易裂果枣品种抗氧化酶活性显著低于抗裂果枣品种[28]。丁改秀等[9]发现,随着枣成熟度的增加,枣果皮的CAT、SOD 活性逐渐降低,表明抗氧化酶活性可能与枣果皮衰老、死亡具有联系,这一研究与本试验中抗氧化酶活性随成熟度增加而降低的趋势一致,喷施BR延缓了果实抗氧化酶活性的下降速度,达到减少裂果的目的。POD 可以消解植物体内生长素,增加木质化程度,POD 活性增强可加速木质化形成,使得果皮细胞死亡,增加裂果风险[29],在本试验中,POD 活性在裂果关键期大量增强,与裂果率趋势一致。

纤维素酶和果胶酶能分解果皮中的纤维素及果胶物质,降低果皮的机械强度,提高果实裂果率[11]。对不同裂性的枣品种纤维素酶活性测定发现,纤维素酶活性随果实成熟度的增加呈先上升后下降趋势,且抗裂枣木枣纤维素酶活性低于易裂枣团枣[30]。BR可以延缓纤维素和果胶含量的减少,并可以抑制纤维素酶和果胶酶的活性[31]。在本试验中,自裂果关键期后,纤维素酶和果胶酶活性呈下降趋势,且BR处理酶活性整体低于对照,这表明外源油菜素内酯降低纤维素酶和果胶酶活性,使其对细胞壁物质的分解速率降低,从而减少了裂果。

3.4 油菜素内酯对骏枣果皮内源激素含量的影响

内源激素的变化是影响果实裂果的重要原因之一,稳定的激素平衡对果实发育及裂果具有重要影响[11]。促生长型激素比值越高,裂果发生的可能性越小[32]。GA3及IAA 是促生长型激素,会影响果实细胞壁的组成和结构,促进细胞壁物质的合成,增加果皮的刚性,有助于维持果实的膨压平衡,有效降低裂果率。ABA是抑制型激素,与GA3之间具有拮抗作用,随着果实成熟度的增加整体呈上升的趋势,ABA 会降解果胶等物质,降低果皮的韧性[33],从而增加果实裂果的风险。本试验结果表明,喷施BR后,GA3含量显著高于对照,ABA 含量除转色期外,各处理皆显著低于对照。全红期各处理浓度(GA3+IAA)/ABA 的比值皆高于对照,与李三玉等[32]的研究结果一致。综上,外源油菜素内酯会影响内源激素含量,从而达到调控枣果实裂果的目的。

4 结 论

喷施油菜素内酯可以有效降低骏枣果实裂果率,提高果实品质、果实硬度、胶黏性、咀嚼性,显著提高果皮CAT和SOD活性及GA3和IAA含量,降低果皮中纤维素酶和果胶酶活性及ABA含量。综上,外源施用油菜素内酯可以降低骏枣果实裂果率,且喷施质量浓度为1 mg·L-1 效果最佳。

参考文献 References:

[1] 李晓鹏,张永凯,李新岗.中国枣产区迁移及驱动力分析[J].西北林学院学报,2024,39(4):267-275.LI Xiaopeng,ZHANG Yongkai,LI Xingang.Analysis on migration and driving forces of Chinese jujube production areas[J].Journal of Northwest Forestry University,2024,39(4):267-275.

[2] BAKEER S M. Effect of ammonium nitrate fertilizer and calcium chloride foliar spray on fruit cracking and sunburn of manfalouty pomegranate trees[J]. Scientia Horticulturae,2016,209:300-308.

[3] 陈斌,吴震,文军琴,林昊维,于璐,薛灵姿,周蓉,蒋芳玲.番茄不规则裂果性状的QTL 定位及候选基因分析[J].园艺学报,2021,48(7):1329-1339.CHEN Bin,WU Zhen,WEN Junqin,LIN Haowei,YU Lu,XUE Lingzi,ZHOU Rong,JIANG Fangling. QTL mapping and candidate genes analysis of irregular fruit cracking in tomato[J].Acta Horticulturae Sinica,2021,48(7):1329-1339.

[4] 王旭旭,樊秀彩,李傲,张超博,房经贵,刘崇怀,上官凌飞.葡萄品种资源裂果性状调查与分析[J].园艺学报,2016,43(11):2099-2108.WANG Xuxu,FAN Xiucai,LI Ao,ZHANG Chaobo,FANG Jinggui,LIU Chonghuai,SHANGGUAN Lingfei. Investigation and analysis on cracking trait in grape berry[J]. Acta Horticulturae Sinica,2016,43(11):2099-2108.

[5] 王振磊,吴巍,王小燕,林敏娟.枣抗裂种质资源筛选与裂果机理初步研究[J].塔里木大学学报,2016,28(3):88-95.WANG Zhenlei,WU Wei,WANG Xiaoyan,LIN Minjuan.Study on pollen germination and pollen tube growth of different cultivars of Ziziphus jujube[J]. Journal of Tarim University,2016,28(3):88-95.

[6] 寸丽芳,房立媛,林敏娟,王振磊.喷施外源钙对骏枣裂果和相关生理特性及显微结构的影响[J]. 果树学报,2023,40(9):1894-1903.CUN Lifang,FANG Liyuan,LIN Minjuan,WANG Zhenlei. Effects of spraying calcium on fruit cracking and related physiological characteristics and microstructure of Junzao jujube[J]. Journal of Fruit Science,2023,40(9):1894-1903.

[7] 杨俊强,王宝明,王小原.枣裂果研究进展[J].山西农业科学,2009,37(3):86-89.YANG Junqiang,WANG Baoming,WANG Xiaoyuan. Research progress of fruit cracking in Chinese jujube[J]. Journal of Shanxi Agricultural Sciences,2009,37(3):86-89.

[8] GHOSH T,PANJA P,SAU S,DATTA P.Role of brassinolide in fruit growth,development,quality and cracking of Litchi cv.Bombai grown in new alluvial zone of west Bengal[J]. International Journal of Bio-Resource and Stress Management,2022,13(5):507-512.

[9] 丁改秀,王保明,王小原,许建庆,陈丽,乔慧琴,仓国营,牛兴艳,温鹏飞.枣果实不同发育期果皮抗氧化酶类活性及MDA和脯氨酸含量变化研究[J].山西农业科学,2014,42(10):1075-1077.DING Gaixiu,WANG Baoming,WANG Xiaoyuan,XU Jianqing,CHEN Li,QIAO Huiqin,CANG Guoying,NIU Xingyan,WEN Pengfei.Studies on the changes of proline,MDA content and activity of POD,SOD and CAT in pericarp during huping jujube development[J]. Journal of Shanxi Agricultural Sciences,2014,42(10):1075-1077.

[10] 李建国,黄旭明,黄辉白.裂果易发性不同的荔枝品种果皮中细胞壁代谢酶活性的比较[J].植物生理与分子生物学学报,2003,29(2):141-146.LI Jianguo,HUANG Xuming,HUANG Huibai. Comparison of the activities of enzymes related to cell-wall metabolism in pericarp between litchi cultivars susceptible and resistant to fruit cracking[J]. Acta Photophysiologica Sinica,2003,29(2):141-146.

[11] 曹一博,李长江,孙帆,张凌云.抗裂与易裂枣内源激素含量和细胞壁代谢相关酶活性比较[J].园艺学报,2014,41(1):139-148.CAO Yibo,LI Changjiang,SUN Fan,ZHANG Lingyun. Comparison of the endogenous hormones content and the activities of enzymes related to cell-wall metabolism between jujube cultivars susceptible and resistant to fruit cracking[J].Acta Horticulturae Sinica,2014,41(1):139-148.

[12] SEYMOUR G B,MANNING K,ERIKSSON E M,POPOVICH A H,KING G J.Genetic identification and genomic organization of factors affecting fruit texture[J].Journal of Experimental Botany,2002,53(377):2065-2071.

[13] 杨植,董梦怡,王振磊,闫芬芬,吴翠云,王玖瑞,刘孟军,林敏娟.基于TPA 法枣酸枣杂交F1 果实质地与裂果对比分析[J].新疆农业科学,2023,60(3):608-615.YANG Zhi,DONG Mengyi,WANG Zhenlei,YAN Fenfen,WU Cuiyun,WANG Jiurui,LIU Mengjun,LIN Minjuan. TPA method was used to analyze the texture and split of F1 fruit of jujube hybrid[J]. Xinjiang Agricultural Sciences,2023,60(3):608-615.

[14] YAMAGUCHI M,SATO I,ISHIGURO M.Influences of epidermal cell sizes and flesh firmness on cracking susceptibility in sweet cherry (Prunus avium L.) cultivars and selections[J].Engei Gakkai Zasshi,2002,71(6):738-746.

[15] CORREIA S,SCHOUTEN R,SILVA A P,GONÇALVES B.Sweet cherry fruit cracking mechanisms and prevention strategies:A review[J].Scientia Horticulturae,2018,240:369-377.

[16] 张川,王亚晨,崔守尧,杨泽恩,吴震,蒋芳玲.耐裂果与易裂果番茄果实发育过程中果实组织衰老与裂果的关系[J].南京农业大学学报,2016,39(4):534-542.ZHANG Chuan,WANG Yachen,CUI Shouyao,YANG Zeen,WU Zhen,JIANG Fangling. The relationship between fruit tissue senescence and fruit cracking in cracking-resistant and susceptible tomato during fruit ripening[J].Journal of Nanjing Agricultural University,2016,39(4):534-542.

[17] 陈光辉,高艳,陈秀娟,谢丽琼.植物激素在植物细胞壁扩展中的作用[J].生命的化学,2012,32(5):464-470.CHEN Guanghui,GAO Yan,CHEN Xiujuan,XIE Liqiong.The role of phytohormones in plant cell wall expansion[J]. Chemistry of Life,2012,32(5):464-470.

[18] 杨帆.不同外源物质处理对番茄裂果及相关性状的影响[D].南京:南京农业大学,2022.YANG Fan. Effects of different exogenous substances on fruit cracking and related traits in tomato[D]. Nanjing:Nanjing Agricultural University,2022.

[19] 王建宇,高秋玲,王振磊,林敏娟.细胞代谢酶活性、碳水化合物及内源激素与枣裂果关系[J].新疆农业科学,2020,57(9):1689-1696.WANG Jianyu,GAO Qiuling,WANG Zhenlei,LIN Minjuan.Relationship between cell metabolism enzyme activity,carbohydrate,endogenous hormones and fruit cracking[J]. Xinjiang Agricultural Sciences,2020,57(9):1689-1696.

[20] 张晶晶.CaCl2处理对‘伏脆蜜’枣裂果生理特性及基因表达与代谢物的影响[D].阿拉尔:塔里木大学,2022.ZHANG Jingjing. Effects of CaCl2 treatment on physiological characteristics,gene expression and metabolites of‘Fucuimi’jujube fruit cracking[D].Ala’er:Tarim University,2022.

[21] 杨途熙,魏安智,郑元,杨恒,杨向娜,张睿.高效液相色谱法同时分离测定仁用杏花芽中8 种植物激素[J].分析化学,2007,35(9):1359-1361.YANG Tuxi,WEI Anzhi,ZHENG Yuan,YANG Heng,YANG Xiangna,ZHANG Rui.Simultaneous determination of 8 endogenous hormones in apricot floral bud by high performance liquid chromatography[J]. Chinese Journal of Analytical Chemistry,2007,35(9):1359-1361.

[22] 侯雷平,李梅兰.油菜素内酯(BR)促进植物生长机理研究进展[J].植物学通报,2001,36(5):560-566.HOU Leiping,LI Meilan. Progress of studies on the plant growth promoting mechanism of brassinolide (BR)[J]. Chinese Bulletin of Botany,2001,36(5):560-566.

[23] 于继洲,马丽萍,张秀梅,范晓峰,王小鸢.枣树裂果机理研究[J].山西农业科学,2002,30(1):76-79.YU Jizhou,MA Liping,ZHANG Xiumei,FAN Xiaofeng,WANG Xiaoyuan. Mechanism of fruit cracking in Chinese jujube[J].Journal of Shanxi Agricultural Sciences,2002,30(1):76-79.

[24] 张鹏飞,高美英,纪薇,牛铁泉,刘亚令.叶片和果实吸水力对枣裂果的影响研究[J].核农学报,2014,28(12):2269-2274.ZHANG Pengfei,GAO Meiying,JI Wei,NIU Tiequan,LIU Yaling. Effects of leaf and fruit water absorbing on fruit cracking in Chinese jujube[J].Journal of Nuclear Agricultural Sciences,2014,28(12):2269-2274.

[25] 贺琰.外源油菜素内酯处理对高温胁迫下赤霞珠葡萄光合特性及果实品质的影响[D].银川:宁夏大学,2022.HE Yan. Effect of exogenous brassinolides treatment on photosynthetic characteristics and fruit quality of Cabernet Sauvignon grapes under heat stress[D].Yinchuan:Ningxia University,2022.

[26] 林竹,苏淑钗,马超,白倩,杨少燕.油菜素内酯对肥城桃光合作用和果实品质的影响[J].经济林研究,2016,34(2):73-78.LIN Zhu,SU Shuchai,MA Chao,BAI Qian,YANG Shaoyan.Effects of EBR on photosynthesis and fruit quality of Feicheng peach[J].Non-wood Forest Research,2016,34(2):73-78.

[27] 院钦,杨四钰,乔帅,任玉锋,周军,张欣,陈卫军,万仲武,魏天军,徐文娣,王惠冉.‘灵武长枣’果实发育过程中果肉质地与解剖结构的变化[J].西北农业学报,2024,33(5):851-861.YUAN Qin,YANG Siyu,QIAO Shuai,REN Yufeng,ZHOU Jun,ZHANG Xin,CHEN Weijun,WAN Zhongwu,WEI Tianjun,XU Wendi,WANG Huiran.Changes in pulp texture and anatomical structure of‘Lingwuchangzao’(Ziziphus jujuba) during fruit development[J].Acta Agriculturae Boreali-occidentalis Sinica,2024,33(5):851-861.

[28] 刘欢,林敏娟,高疆生,王振磊.枣果皮细胞壁代谢酶活性与抗裂果的关系[J].新疆农业科学,2018,55(5):845-854.LIU Huan,LIN Minjuan,GAO Jiangsheng,WANG Zhenlei.Study on the relationship between the activity of cell wall metabolic enzymes and anti-cracked in jujube peel[J].Xinjiang Agricultural Sciences,2018,55(5):845-854.

[29] 徐如涓,李向东,何宇炯,王玉琴,赵毓橘.表油菜素内酯和胆甾内酯对葡萄座果和成熟的影响[J].上海农学院学报,1994,12(2):90-95.XU Rujuan,LI Xiangdong,HE Yujiong,WANG Yuqin,ZHAO Yuju.Effects of treatments with epibrassinolide and chololic lactone on the fruit-set and ripening in some grape cultivation[J].Journal of Shanghai Agricultural College,1994,12(2):90-95.

[30] 栗现芳,姚瑞,赵瑞华,陈国梁,陈宗礼.枣果纤维素酶对裂果发生的影响[J].安徽农业科学,2016,44(27):65-68.LI Xianfang,YAO Rui,ZHAO Ruihua,CHEN Guoliang,CHEN Zongli. Effect of cellulase on jujube fruit cracking[J].Journal of Anhui Agricultural Sciences,2016,44(27):65-68.

[31] ZENG Y Y,ZHANG J M,SU H,XIE L,ZHAO Y,XIONG Y B,ZHOU W H. Cell wall and reactive oxygen metabolism responses of stored Shine Muscat grapes to combined melatonin and 24-epibrassinolide treatment[J]. Postharvest Biology and Technology,2025,219:113192.

[32] 李三玉,陈苑虹,吕均良,林颖.玉环柚果实内源激素含量与裂果关系的研究[J].科技通报,1999,15(3):166-169.LI Sanyu,CHEN Yuanhong,LÜ Junliang,LIN Ying. Studies on the relationship between fruit hormones content and fruit cracking in Yuhuan pomelo[J]. Bulletin of Science and Technology,1999,15(3):166-169.

[33] 卢文静.脱落酸和生长素调控香蕉及草莓果实成熟的作用机理[D].杭州:浙江大学,2018.LU Wenjing. Regulation of abscisic acid and indole-3-acetic acid on ripening of banana and strawberry fruit[D]. Hangzhou:Zhejiang University,2018.

Effect of exogenous Brassinolide on improving fruit quality and reducing fruit cracking in Junzao jujube

YANG Cheng,WANG Zhenlei,ZHANG Chuanjiang,LIU Wei,WU Cuiyun,LIN Minjuan*
(College of Horticulture and Forestry of Tarim University/Southern Xinjiang Characteristics of Fruit Trees High Quality Cultivation and Deep Processing Technology of National Local Joint Engineering Laboratory/Tarim Basin Biological Resources Protection and Utilization of the Ministry to Build a National Key Laboratory Cultivation Base/Tarim Oasis Agricultural Key Laboratory of the Ministry of Education,Alar 843300,Xinjiang,China)

Abstracts:【Objective】Xinjiang is the largest jujube producing area in China,and Junzao is one of the main cultivated varieties in Xinjiang. Jujube fruit cracking problem has been plaguing the development of jujube industry, causing serious economic losses. Brassinolide can improve fruit quality, increase fruit hardness, regulate fruit endogenous hormone content and increase peel toughness. Therefore, the purpose of this study was to investigate the effects of different concentrations of brassinolide on fruit cracking,nutritional quality,texture,cell wall metabolic enzyme activity and endogenous hormone content in Junzao.【Methods】In this experiment, 10-year-old trees were used as experimental materials.Different concentrations of brassinolide(1 mg·L-1,2 mg·L-1 and 3 mg·L-1)were sprayed during fruit expansion,and clear water was sprayed as the control.The fruit cracking rate,fruit quality(soluble solids,soluble sugar, titratable acid, hardness, adhesiveness, elasticity, chewiness and adhesiveness), peel cell wall metabolic enzymes (catalase (CAT), peroxidase (POD), superoxide dismutase (SOD), cellulase,pectinase)and peel endogenous hormones(GA3,IAA and ABA)were measured at green-ripening stage,white-ripening stage,color-turning stage,half-red stage and full-red stage.Excel 2016 was used for data collection and analysis, SPSS 27.0 was used for significance analysis, Origin 2021 was used for mapping,and correlation analysis and principal component analysis were carried out for each index at whiteripe stage.【Results】Spraying brassinolide during the fruit enlargement period of Junzao can significantly reduce the fruit cracking rate. The increase of fruit cracking rate with different treatments at white-ripe stage was the largest,which was the key period of fruit cracking.The fruit cracking rate was the highest at the whole-red stage, which was the peak period of fruit cracking.The fruit cracking rate of different brassinolide concentrations was 6.38%, 4.63% and 4.48% lower than that of the control.The soluble solids content in fruits treated with brassinolide increased by 2.49%,5.09%and 33.84%,respectively, and the soluble sugar content increased by 3.35%, 12.72% and 7.08%, respectively, compared with the control. Spraying brassinolide can increase fruit hardness, chewiness and gumminess.The activities of CAT and SOD in each treatment decreased with the increase of maturity.The CAT activity in the peel under different brassinolide concentrations increased by 18.54%,39.07%and 98.01%,respectively, compared with the control. The SOD activity increased by 55.32%, 67.25% and 71.18%,respectively, and the POD activity decreased by 24.66%, 1.15% and 32.06%, respectively, compared with the control. The activities of cellulase and pectinase reached the peak at the white-ripening stage,and then decreased with the increase of fruit maturity.The activities of pectinase at the whole-red stage with each treatment were 6.92%, 7.75% and 13.91% lower than those of the control, respectively. The content of endogenous hormones in peel changed significantly with fruit growth and development.GA3 and IAA showed a downward trend, and GA3 and IAA in the peel of fruits treated with brassinolide were significantly or extremely significantly higher than those in the control at each stage. Gibberellin GA3 increased by 8.30%, 24.70% and 14.07%, respectively, at the whole-red stage compared with the control.IAA content increased by 312.12%,198.48%and 457.58%,respectively.The ABA content was significantly reduced by 16.30%, 22.47% and 17.10%, respectively, compared with the control. Hormone balance is one of the factors affecting fruit cracking. At the full-red stage of fruit, the ratio of(GA3+IAA)/ABA in the pericarp of fruits treated with brassinolide and water(control)was 49∶57∶55∶33,respectively,which was higher than that of the control.The correlation between different concentrations of brassinolide treatment and control at white-ripe stage was analyzed. The analysis showed that the fruit cracking of jujube fruit was related to many factors such as cell wall metabolic enzymes, fruit quality and endogenous hormones.Fruit cracking rate was significantly and positively correlated with titratable acid content.There was a significant positive correlation with ABA content.It was significantly and negatively correlated with hardness, adhesion, CAT activity and GA3 content. There was a significant negative correlation with SOD activity. Principal component analysis showed that the main traits affecting fruit quality were hardness, chewiness, adhesiveness, SOD activity, GA3 content and fruit cracking rate.【Conclusion】Spraying brassinolide can effectively reduce the fruit cracking rate of Junzao,and improve fruit soluble solids content,soluble sugar content,fruit hardness,chewiness and adhesiveness.The CAT and SOD activities as well as GA3 and IAA contents in the pee increased,and the cellulase and pectinase activity as well as ABA content decreased. The cracking resistance was improved and the cracking rate was reduced.Among them, spraying 1 mg·L-1 concentration of brassinolide had the best effect on preventing fruit cracking.

Key words: Junzao jujube;Brassinolide;Fruit cracking;Antioxidant enzyme activity

中图分类号:S665.1

文献标志码:A

文章编号:1009-9980(2025)05-1010-13

DOI: 10.13925/j.cnki.gsxb.20240661

收稿日期:2024-11-19

接受日期:2025-02-21

基金项目:国家自然科学基金项目(31560541);兵团中青年科技创新领军人才计划(2018CB032);兵团重点领域科技攻关计划(2023AB004-04)

作者简介:杨成,男,在读硕士研究生,主要从事果树种质资源筛选的研究。E-mail:yangc0226@163.com

*通信作者 Author for correspondence.E-mail:lmjzky@163.com