基于越橘南高丛与北高丛正反交群体的果实贮藏相关性状筛选及其遗传倾向

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刘有春^,¹^,², 刘威生¹, 王兴东¹, 杨艳敏¹, 魏鑫¹, 孙斌¹, 张舵¹, 杨玉春¹, 刘成^,¹, 李天忠^,²

¹辽宁省果树科学研究所,辽宁营口 115009

²中国农业大学园艺学院,北京 100193

Screening and Inheritance of Fruit Storage-Related Traits Based on Reciprocal Cross of Southern×Northern High Bush Blueberry (Vaccinium Linn)

LIU YouChun^,¹^,², LIU WeiSheng¹, WANG XingDong¹, YANG YanMin¹, WEI Xin¹, SUN Bin¹, ZHANG Duo¹, YANG YuChun¹, LIU Cheng^,¹, LI TianZhong^,²

¹Liaoning Institute of Pomology, Yingkou 115009, Liaoning

²College of Horticulture, China Agricultural University, Beijing 100193

通讯作者: 刘成,E-mail： stevecliu@hotmail.com; 李天忠,E-mail： litianzhong1535@163.com

责任编辑: 赵伶俐
收稿日期:2020-02-24接受日期:2020-04-21网络出版日期:2020-10-01

基金资助:

国家自然科学基金青年基金.31701881
辽宁省重点研发计划.2019JH2/10200010
辽宁省农业科学院学科建设计划.2019DD164924
地方专业性技术创新平台.2018103002

Received:2020-02-24Accepted:2020-04-21Online:2020-10-01
作者简介 About authors
刘有春,Tel：18641713730;E-mail： liuyouchun911@126.com。

摘要
【目的】筛选与越橘果实贮藏性相关的性状并揭示其遗传规律,为越橘果实贮藏性改良育种提供依据。【方法】调查越橘果实的果蒂痕大小、果实硬度、可溶性固形物含量、酸含量、果实重量、果型指数、贮藏性等7个性状,通过相关性分析和主成分分析筛选与贮藏性相关的主要性状,利用正反交群体分析其遗传倾向。【结果】相关性分析和主成分分析结果表明,越橘果实贮藏性与果蒂痕性状呈极显著负相关（P<0.001）,与果实硬度性状呈极显著正相关（P<0.01）,二者是与蓝莓贮藏性相关的主要性状。越橘果实带柄和脱柄贮藏试验表明果蒂痕大小对越橘果实贮藏有重要影响。果蒂痕大小和果实硬度性状在正、反交群体中呈正态分布,表现为多基因控制的数量性状,变异系数均大于15%。正、反交群体果蒂痕大小趋近于亲本平均值,无父母本倾向,其广义遗传力（H²）在正、反交群体中分别为0.887和0.867,其遗传变异主要来自遗传效应,遗传潜能较大。正、反交群体果实硬度平均值均趋于低值亲本,即存在倾向于果实硬度变小的劣质遗传,H²在正、反交组合中分别为0.535和0.514,表明越橘果实硬度遗传效应较小,遗传不稳定,易受环境影响。【结论】越橘果实果蒂痕大小和果实硬度性状与贮藏性密切相关,其中果蒂痕性状呈趋中遗传,其遗传变异主要来自遗传效应,遗传潜能大,超高亲遗传不明显;果实硬度性状遗传受低值亲本影响较大,存在明显的劣质遗传倾向,遗传效应小,受环境影响大。
关键词： 越橘;贮藏性;果蒂痕;果实硬度;正反交;广义遗传力

Abstract

【Objective】 This study was aimed to screen the diagnostic phenotypic traits for the storability of blueberry fruit and analyze their inheritance to assist the storage improvement breeding. 【Method】 Seven phenotypic traits were investigated, including stem scar, berry firmness, soluble solids content, acid content, fruit weight, fruit index and storability, among which the storability related traits was screened through correlation analysis and principal component analysis (PCA), and the inheritance was analyzed with reciprocal cross progenies.【Result】 The correlation analysis and PCA showed that stem scar was negatively correlated with fruit storability (P<0.001), but berry firmness was positively correlated with fruit storability (P<0.01). Both stem scar and berry firmness played major roles in the development of the storability of blueberry fruit. The storage test of fruit with and without stalk revealed that stem scar had a great influence on the storability of blueberry. The size of stem scar and berry firmness presented normal distributions in reciprocal populations with the coefficient of variation of more than 15%, suggesting that stem scar and berry firmness were quantitatively controlled by polygenes. The mean of stem scar in the reciprocal populations was similar to the mid-parental values, and its broad sense heritability (H²) in reciprocal populations were 0.887 and 0.867, respectively, indicating that the variations of stem scar mainly resulted from inheritance, and were mainly affected by additive effects. The mean of berry firmness in both reciprocal populations was prone to low value parent, and its broad sense heritability was 0.585 and 0.514, respectively in reciprocal populations, indicating that berry firmness was affected to a great extent by environmental effects. 【Conclusion】 The stem scar and berry firmness were the major traits for blueberry fruit storability. The inheritance of stem scar inclined to the mid-parental value, and its variation was mainly controlled with genetic effects, while berry firmness value tended to the low-value parent with large environmental effects.

Keywords：blueberry;storability;stem scar;berry firmness;reciprocal population;genetic heritability

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本文引用格式
刘有春, 刘威生, 王兴东, 杨艳敏, 魏鑫, 孙斌, 张舵, 杨玉春, 刘成, 李天忠. 基于越橘南高丛与北高丛正反交群体的果实贮藏相关性状筛选及其遗传倾向[J]. 中国农业科学, 2020, 53(19): 4045-4056 doi:10.3864/j.issn.0578-1752.2020.19.017
LIU YouChun, LIU WeiSheng, WANG XingDong, YANG YanMin, WEI Xin, SUN Bin, ZHANG Duo, YANG YuChun, LIU Cheng, LI TianZhong. Screening and Inheritance of Fruit Storage-Related Traits Based on Reciprocal Cross of Southern×Northern High Bush Blueberry (Vaccinium Linn)[J]. Scientia Acricultura Sinica, 2020, 53(19): 4045-4056 doi:10.3864/j.issn.0578-1752.2020.19.017

0 引言

【研究意义】越橘（Vaccinium corymbosum Linn）,商品名蓝莓（Blueberry）,果实营养丰富、风味独特,备受消费者青睐。然而,越橘果实汁多、皮薄,成熟期主要集中在高温多雨的夏季,采后呼吸作用旺盛,蒂部易受霉菌侵染,导致果实快速软化、褐变并腐烂变质^[1,2,3],极不耐运输和贮藏。货架期短是制约越橘产业发展的瓶颈问题之一。近几年快速发展的保鲜技术通过各种物理、化学保鲜措施可有效延长越橘果实贮藏期。然而,任何保鲜措施都依赖果实自身耐贮性,在遗传水平上解析耐贮性相关性状及其遗传规律,通过育种手段改善果实自身贮藏性,提高越橘果实保鲜效果,对越橘产业发展具有重要意义。【前人研究进展】基于越橘果实不耐贮藏的特性,目前研究大多集中在采后保鲜技术、采后衰变机理、生理代谢等方面。据报道,越橘果实采后预冷措施保鲜期能延长一倍以上^[4],冰温保鲜是越橘果实较理想的保鲜方式,冰温条件下越橘果实呼吸强度降低,失重率、果实硬度、口感及营养等变化减小^[5,6]。此外,利用1-MCP（1-甲基环丙烯）气调处理抑制乙烯产生^[7,8,9,10]、控制呼吸强度等,从而延缓果实后熟软化^[11,12,13];通过保鲜涂膜^{[14,15,16,17]}、植物精油^[18,19,20]、⁶⁰Co-γ辐照^[2,21-22]、熏蒸^[23,24]等措施抑制病原菌活性,从而达到越橘果实贮藏保鲜的目的。目前鲜见越橘果实贮藏相关性状筛选及性状遗传方面的研究报道,耐贮运品种选育工作相对落后。【本研究切入点】目前有关越橘采后贮藏方面的研究忽视了果实本身的贮藏性,在改善果实本身耐贮性的基础上,保鲜措施才能最大程度发挥作用,达到最佳贮藏保鲜效果。因此,通过育种提高、改良越橘果实的贮藏性基础（如果蒂痕、果实硬度等）十分必要。【拟解决的关键问题】本研究通过特定贮藏条件下,筛选与果实贮藏性密切相关的性状,利用正、反交群体反映出性状的遗传倾向和特点,为越橘果实贮藏保鲜和越橘果实贮藏性改良育种提供参考依据。

1 材料与方法

试验于2018—2019年在辽宁省果树科学研究所进行。

1.1 材料

1.1.1 贮藏相关性状筛选的试材试材选择种植于辽宁省果树科学研究所资源圃的26份越橘品种和育种圃的98份杂交后代（表1）,试材模型丰富,可较全面表征果实质量、果型指数、果蒂痕大小、果肉硬度、可溶性固形物含量、酸含量及贮藏性特征。

Table 1
表1
表1供试材料
Table 1Test materials

栽培/杂交组合类型组合 Cultivation/Hybrid cross type	品种份数 Accessions of variety	品种名称 Name of variety
北高丛越橘 V. corymbosum northern highbush (NHB)	12	斯巴坦 Spartan,蓝丰 Bluecrop,伯克利 Berkeley,泽西Jersey,布里吉塔 Brigitta,杜克 Duke,埃利奥特 Elliott,晚蓝 Lateblue,达柔 Darrow,喜来 Sierra,日出 Sunrise,早蓝 Earliblue
南高丛越橘 V. corymbosum southern highbush (SHB)	7	奥尼尔 O'Neal,密斯梯 Misty,佐治亚宝石 Georgiagem,莱格西 Legacy,比洛克西Biloxi,N6,N5
半高丛越橘 V. corymbosum half-highbush	4	北陆 Northland,北村 Northcountry,齐佩瓦 Chippewa,北蓝 Northblue
矮丛越橘 V. augustifolium	1	美登Blomidon
兔眼越橘 V.virgatum	2	园蓝 Gardenblue,梯扶蓝 Tifblue
北高丛×北高丛 NHB×NHB	33	斯巴坦 Spartan×蓝丰 Bluecrop,泽西Jersey×斯巴坦 Spartan,康维尔Coville×斯巴坦 Spartan
北高丛×南高丛 NHB×SHB	29	斯巴坦 Spartan×N6,斯巴坦 Spartan×奥尼尔 O'Neal,伯克利 Berkeley×N6
南高丛×北高丛 SHB×NHB	17	N5×伯克利 Berkeley,N6×斯巴坦 Spartan
南高丛×南高丛 SHB×SHB	19	奥尼尔 O'Neal×密斯梯 Misty

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1.1.2 贮藏相关性状遗传倾向分析的试材

试验材料为南高丛越橘品种‘N6’（V. corymbosum SHB）、北高丛越橘品种‘Berkeley’（V. corymbosum NHB）及其291株杂交后代（已通过真杂种鉴定）。亲本‘N6’具有小果、圆形、果蒂痕中等大小、果肉较软、风味浓郁等特征;‘Berkeley’为大果、扁圆形、果蒂痕大、果实硬度较高、风味偏淡的越橘品种。正交群体‘Berkeley’×‘N6’120株,反交群体‘N6’×‘Berkeley’171株。2008年建立正反交群体,2011年起开花结果,现已全面进入盛果期。杂种苗定植株行距为0.6 m×1.0 m,土壤pH约4.8,生长期间亲本与杂交群体采用相同的施肥、灌溉、修剪等管理措施。

1.2 方法

1.2.1 果实采收采样方法参照刘有春等^[25,26]的方法进行,在树体果实大部分进入成熟期后,于晴天上午8：00—10：00选取树体中上部外围的已具有成熟特征的果实采收,即果实充分膨大,果皮完全转变成紫色或紫黑色,果蒂粉色退净,充分表现品种的固有风味。为尽量保证亲本及后代果实成熟度的一致性,取样由同一实验者完成,采后于-50℃下保存备用。

1.2.2 果实表型性状数据收集平均单果重采用称重法测量;果型指数=纵径/横径,纵径和横径采用数显游标卡尺测量;果蒂痕大小为数显游标卡尺测量的果蒂痕直径（mm）;可溶性固形物与总酸含量采用越橘专用数显糖酸一体机（ATAGO,日本）测定;果肉硬度采用浆果专用便携式硬度计测定。以上指标均为3次重复,每次重复测量10个果实,取平均值用于后续分析。

1.2.3 贮藏期计算在供试越橘果实贮藏试验中,通过恒温冰箱（海尔）设定温度4℃,空气湿度60%,降低外界因素变化影响贮藏室环境。每份样品取100粒越橘果实装入容器（带孔）,每隔5 d调查一次,当10%样品出现皱缩/发霉/软化则视为样品贮藏期结束,并记录贮藏天数。

1.2.4 贮藏性对比试验处理为了验证果蒂痕对越橘果实贮藏的重要性,越橘果实设果蒂痕小于1.0 mm（A）、1.1—2.0 mm（B）、2.1—3.0 mm（C）、大于3.0 mm（D）4个处理,进行相同条件下（恒温4℃）带柄贮藏（模拟果蒂无伤口）和脱柄贮藏（常规方式）对比试验,每处理150 g,每5 d调查1次,测量果实硬度变化,拍照记录果实状态。

1.3 数据统计分析

1.3.1 遗传相关性及主成分分析 7个供试性状的表型数据,取其平均值（±标准差）用于统计分析。采用“Pearson”方法计算相关性（correlation）和显著性差异分析;随后通过离差标准化法对数据进行标准化转化为无量纲数值,经“factoextra”软件包进行主成分分析（principal component analysis,PCA）,以上分析均在R中进行。

1.3.2 遗传分析根据CHEN等^[27]报道的方法计算正反交群体的广义遗传力（The broad sense heritability,H²）,根据崔艳波等^[28]的方法计算变异系数（coefficient of variation,CV）,参照刘有春等^[29]的方法计算超亲遗传,包括超高亲（Ratio of higher than high parent,HH）、超中亲（Ratio of higher than middle parent,HM）、超低亲（Ratio of lower than low parent,LL）。上述分析均通过R中“ggplots 2”^[30]分析包完成绘图。

2 结果

2.1 越橘果实性状分布范围分析

124个供试品种/杂交后代果实的单果重、果蒂痕大小、果形指数、果实硬度、可溶性固形物、总酸和贮藏性性状的范围值如图1所示。多数样品单果重分布范围在0.70—3.11 g,平均2.00 g,1个样品单果重大于5.00 g;果蒂痕直径分布范围在0.75 mm—4.29 mm,平均2.55 mm;果形指数分布范围相对较小,在0.67—0.94,1个样品果形指数1.02,为圆形果实;果实硬度的分布范围在0.12—0.30,有2个样品硬度值大于0.30,分别为0.33和0.39,明显高于其他供试样品;可溶性固形物含量分布范围在9.32%—16.67%,其中含量值大于13%的样品占36.94%;总酸含量分布范围在0.14%—0.80%,有2个样品的总酸含量大于1.00%,分别是1.02%和1.15%,口感酸度大;果实贮藏期（4℃）在9—38 d,有2个样品贮藏期大于40 d,分别为40和46 d,为耐贮品种,贮藏期≥30 d的品种/杂交后代占供试样品总数的22.3%。

图1

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图1供试样品果实7个性状数量值分布范围

箱体高度等于四分位间距,代表数据的集中分布范围,包含样本50%的测定数据,箱体中间横线表示中间值,上下截止线之间包含了99%的样本测定数据,星号（*）表示超出本体值外的极值
Fig. 1A range of seven traits of blueberry fruit tested

The height in a box is equal to the interquartile distance,indicating the distribution for 50% of the data, the horizontal lines in the interior of each box are the median values. Approximately 99% of the data falls inside the whiskers (the dotted lines extending from the top and bottom of the box). The data outside these whiskers are indicated by asterisk (*)

2.2 相关性分析与越橘果实贮藏相关性状筛选

采用二项正态分布置信检测方法对每一项指标进行两两置信检测,采用Pearson相关系数分析供试性状间的相关性,筛选与越橘果实贮藏性相关性状,（表2）。单果重与果型指数呈极显著负相关（P<0.001）,与可溶性固形物含量呈显著负相关（P<0.05）,表明单果重越大,果实性状越趋近于扁圆形,反之则趋近于圆形或近圆形,且单果重增加会影响可溶性固形物含量的积累。贮藏性与果蒂痕大小呈极显著负相关（P<0.001）,与果实硬度呈极显著正相关（P<0.01）,表明果蒂痕大严重影响越橘果实贮藏,而果实硬度大则有利于越橘果实贮藏。

Table 2
表2
表2越橘果实供试性状相关性分析
Table 2Correlation analysis among tested traits of blueberry fruit

指标 Index	单果重 Fruit weight	果蒂痕 Stem scar	果型指数 Index	果实硬度 Berry firmness	可溶性固形物含量 Soluble solids content	酸含量 Acid content	贮藏性 Storability
单果重 Fruit weight	1
果蒂痕 Stem scar	0.1	1
果型指数 Index	-0.724***	-0.136	1
果实硬度 Berry firmness	0.147	-0.146	-0.105	1
可溶性固形物含量 Soluble solids content	-0.227*	0.113	0.142	-0.034	1
酸含量 Acid content	-0.201	-0.167	0.245*	0.077	0.105	1
贮藏性 Storability	-0.053	-0.527***	-0.03	0.315**	0.145	0.118	1

*, ** and *** means significant at 0.05, 0.01and 0.001 level, respectively
*、**和***分别表示在0.05、0.01和0.001水平上显著相关

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上述相关性分析表明果蒂痕与果实贮藏时间极显著相关（P<0.001）,以下通过带柄贮藏（模拟果蒂无伤口）和脱柄贮藏（常规方式）对比进一步验证果蒂痕对越橘果实贮藏的影响。如图2所示,果蒂痕4个处理在4℃恒温下贮藏20 d后,带柄贮藏（左）越橘果实表观新鲜度明显好于脱柄贮藏（右）,脱柄贮藏处理随果蒂痕增大,果实皱缩及发霉程度增加,而带柄贮藏各处理果实表观均新鲜,无明显差异。如表3所示,带柄贮藏20 d后,4个处理越橘果实硬度稍有降低,降低幅度分别为9.25%、13.28%、13.79%和14.81%,而脱柄处理果实硬度快速降低,降低幅度分别为57.27%、59.77%、65.95%和71.76%,表明果蒂痕是影响越橘果实贮藏时间的主要因素之一。

图2

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图2果蒂痕对越橘果实贮藏性的影响（4℃贮藏20 d）

A—D表示果蒂痕大小。A：≤1.0 mm,B：1.1—2.0 mm,C：2.1—3.0,D：>3.0 mm;左：带柄贮藏,右：不带柄储藏
Fig. 2Effect of stem scar size on blueberry storability（4℃ storage for 20 days）

A-D represent stem scar size. A: ≤1.0 mm, B:1.1—2.0 mm, C:2.1—3.0, D: >3.0 mm. Left: storage with fruit stalks, Right: storage without fruit stalks

Table 3
表3
表34℃贮藏过程中越橘果实硬度变化
Table 3Firmness change of blueberry fruit during 4℃ storage

处理 Treatment	贮藏前果实硬度 Berry firmness before storage (kg?cm^-2)	带柄贮藏 Storage with fruit stalks			脱柄贮藏 Storage without fruit stalks
处理 Treatment	贮藏前果实硬度 Berry firmness before storage (kg?cm^-2)	20 d后果实硬度 Berry firmness of after 20 days (kg?cm^-2)	降低幅度 Reduced extent of berry firmness (%)	新鲜/皱缩/发霉 Fresh/shrunk/ moldy	20 d后果实硬度 Berry firmness of after 20 days (kg?cm^-2)	降低幅度 Reduced extent of berry firmness (%)	新鲜/皱缩/发霉 Fresh/shrunk/ moldy
A	0.227	0.206	9.25	新鲜 Fresh	0.097	57.27	皱缩 Shrunk
B	0.256	0.222	13.28	新鲜 Fresh	0.103	59.77	皱缩 Shrunk
C	0.232	0.200	13.79	新鲜 Fresh	0.079	65.95	皱缩 Shrunk
D	0.206	0.184	14.81	新鲜 Fresh	0.061	71.76	皱缩+发霉 Shrunk+Moldy

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2.3 越橘果实贮藏性相关性状主成分分析

对124个供试越橘品种/品系的果实性状进行主成分分析,将单果重、果蒂痕大小、果实硬度等7个性状转化为7个主成分,根据各主成分的特征值和贡献率进行主成分提取,如表4所示,前4个主成分累计方差贡献率为79.828%,反映了性状的主要信息。其中第一主成分和第二主成分方差贡献率分别是28.583%和24.159%,为最主要主成分,第三主成分和第四主成分方差贡献率分别是14.610%和12.476%。为了更好地解释各指标与主成分因子之间的对应关系,绘制平面图,以载荷值大小反映各变量在主成分中的重要程度（图3）。第一主成分（PC1）方差贡献率为28.583%,与果型指数、酸含量、贮藏时间呈正相关,与果实重量和果蒂痕大小呈负相关,其中果型指数和果实大小的方差占PC1方差的66.494%,即主要代表了外观性状的方差,果型指数和果实大小呈负相关。第二主成分（PC2）方差贡献率为24.159%,与果实贮藏时间、果实硬度、果实大小正相关,与果蒂痕大小、果型指数负相关,其中果实贮藏时间、果蒂痕大小和果实硬度的方差占PC2方差的84.324%,即主要代表了贮藏相关性状的方差,可以看出果实贮藏时间与果肉硬度呈正相关,与果蒂痕大小呈负相关。

Table 4
表4
表4越橘果实供试性状主成分值
Table 4PCs scores of blueberry tested traits

主成分 Principal component	特征值 Eigenvalue	方差贡献率 Variance contribution rate (%)	累计方差贡献率 Cumulative variance contribution rate (%)
1	2.001	28.583	28.583
2	1.691	24.159	52.742
3	1.023	14.610	67.352
4	0.873	12.476	79.828

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图3

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图3越橘果实供试性状第一、第二主成分平面图

括号中百分数代表各主成分所占的方差贡献率
Fig. 3Plots of blueberry tested trait for the first two PCs

The percentage in brackets represents the variance contribution rate of each principal component

2.4 越橘果实贮藏主要相关性状的遗传变异

相关性分析、果实贮藏对比和主成分分析表明果实蒂痕大小和果实硬度是影响越橘贮藏性的主要因素,以下对这两个性状进行遗传趋势分析。

如表5所示,亲本‘N6’和‘Berkeley’果蒂痕大小和果实硬度性状存在差异,‘N6’果蒂痕较小（1.964 mm）且果实硬度小（0.189 kg?cm^-2）; ‘Berkeley’果实硬度大（0.278 kg?cm^-2）且果蒂痕大（3.501 mm）。二者组配杂交群体,果蒂痕大小和果实硬度性状在正、反交后代中均广泛分离,变异系数均在15%以上。正、反交群体后代果蒂痕大小平均值分别是2.847和2.683 mm,无明显差异,趋近于亲本平均值2.732 mm,无父母本倾向。果蒂痕大小的广义遗传力（H²）在正、反交群体中分别是0.887和0.867,表明性状的变异主要来自遗传效应,遗传潜能大。正、反交群体后代果实硬度平均值分别为0.185和0.186 mm,无明显差异,趋近于低值亲本,即杂交后代倾向于果实硬度变小的劣质遗传。果肉硬度广义遗传力（H²）在正、反交群体中分别是0.535和0.514,表明越橘果实硬度遗传效应小,遗传不稳定,易受环境影响。

Table 5
表5
表5杂交后代果蒂痕大小和果实硬度的广义遗传力和超亲遗传
Table 5Broad-sense heritability and transgressive inheritance of stem scar size and berry firmness in progeny populations

性状 Trait	亲本 Parent			正交群体 N6×Berkeley							反交群体 Berkeley×N6
性状 Trait	N6	Berkeley	?P	?x±S	范围Range	CV (%)	H²	HM (%)	HH (%)	LL (%)	?x±S	范围 Range	CV (%)	H²	HM (%)	HH (%)	LL (%)
果蒂痕大小 Stem scar size (mm)	1.964	3.501	2.732	2.847±0.534	1.412– 4.29	18.750	0.887	48.538	5.848	9.357	2.683±0.579	0.750–4.018	21.590	0.867	61.667	10.000	6.667
果实硬度 Berry firmness (kg?cm^-2)	0.189	0.278	0.234	0.185±0.035	0.080– 0.27	18.900	0.535	10.526	1.754	53.801	0.186±0.038	0.086– 0.282	20.240	0.514	9.167	0.000	56.667

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从超亲遗传趋势来看,对于果蒂痕超低亲（LL）为优势遗传,正、反交群体的超低亲比率（LL）较低,分别为9.357%和6.667%,而劣质遗传,即超高亲比率（HH）也较低,分别为5.848%和10%,说明越橘果实果蒂痕大小的杂种优势不明显。对于果实硬度而言,超高亲为优势遗传,但在杂交后代中超高亲率均极低,正交群体仅1.754%,反交群体中无果实硬度大于‘Berkeley’（0.278 kg?cm^-2）的后代,超中亲率也仅为9.127,而劣质遗传,即超低亲率在正、反交群体中分别高达53.801%和56.667%,说明越橘果实硬度存在明显的劣质遗传趋势。

如图4-A、C所示,正、反交群体后代的果蒂痕直径多集中在双亲之间,极低或极高值后代仅占少数,频率分布曲线呈单峰正态分布,说明越橘果实果蒂痕直径是多基因控制的数量性状。如图4-B、D所示,正、反交群体后代的果实硬度频率分布曲线基本呈单峰正态分布,多数集中在0.15—0.22 kg?cm^-2,但半数杂交后代集中在低硬度区域,仅分离出极少数硬度值大于或接近高值亲本的后代。

图4

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图4越橘正、反交群体果蒂痕直径和果实硬度频率分布和曲线图

A、B表示 Berkeley×N6群体,C、D表示N6×Berkeley群体;虚线表示群体平均值
Fig. 4Histogram of frequency distribution and probability density curve of stem scar size and berry firmness in blueberry reciprocal populations

A and B represent population of Berkeley×N6, C and D represent population of N6×Berkeley, dotted line represent mean value of population

3 讨论

3.1 越橘果蒂痕和果实硬度性状对果实贮藏性的影响

果蒂痕是果柄与果实脱离时由于果皮破裂和果肉组织损伤而形成的小伤痕,通常是果实受霉菌侵染和失水的关口。葡萄、樱桃、草莓、软枣猕猴桃等果实带柄采收,尤其是葡萄,防止脱粒是提高商品性的重要指标。研究发现葡萄果柄粗细、果蒂痕大小与贮藏性密切相关,‘巨峰’葡萄果柄细,其果蒂面积小,单位面积果柄和果蒂承受的浆果自重大,导致果粒易在运输中脱落,果穗松动导致果蒂与果粒之间产生微小的伤痕,引起霉变进而加重脱粒,而‘秋红’等葡萄果柄粗,果蒂面积较大,耐拉力较大,单位面积果柄和果蒂承受的自重较小,果粒不易脱落^[31]。本研究发现越橘果实果蒂痕对果实贮藏性的方差贡献率最大,其大小与贮藏时间呈极显著负相关,即果蒂痕越小,果实贮藏性越好,反之贮藏性越差,与葡萄有所不同,可能与葡萄和越橘果实的采收方式存在本质区别有关。葡萄整穗采收,所以防止脱粒是商品性、贮藏性的关键,果柄粗、果蒂面积大可减小单位面积果柄和果蒂承受的自重而减少脱粒;但越橘果实脱柄单粒采收,防止果蒂痕处霉变和失水是商品性和贮藏性的关键,果柄与果实脱离时果皮破损面积越小越有利于贮藏。此外,本研究通过越橘果实带柄贮藏和脱柄常规贮藏对比发现,果实带柄贮藏（4℃）20 d后,果实仍保持新鲜、果肉硬度小幅度下降,而脱柄贮藏后果实失水皱缩甚至果蒂痕发霉,果实硬度大幅降低,失水皱缩及硬度降低程度随着果蒂痕增大而增加,进一步证实了果蒂痕是影响越橘果实贮藏的关键因素。所以,在生产中选择果蒂痕小的品种对于提高果实贮藏性具有重要意义。

果实硬度是受果实自身质地、细胞结构、特殊离子成分以及果胶变化、采后呼吸代谢、酶活性、激素调节、温度等多种因素影响的复杂性状^[32,33,34]。研究发现,硬度对葡萄、番茄、草莓果实贮藏性有重要影响,硬度越大贮藏性越好^[34,35,36]。本研究发现,越橘果实硬度与贮藏时间呈显著正相关,对果实贮藏性的方差贡献仅次于果蒂痕大小。与草莓、葡萄等果实一样,越橘果实硬度也是影响越橘果实贮藏的主要因子,在生产上主要通过预冷^[4]、冰温贮藏^[5,6]及1-MCP^[7,8,9,10]、气调箱^[11,12,13]处理,以抑制乙烯产生、控制呼吸强度等,从而延缓果实后熟软化。同时,笔者调查发现,越橘果实硬度在品种间存在差异,果皮厚、果肉致密、种子少的品种果实硬度普遍较高,可见果皮厚度、果肉质地和种子数量等与果实硬度相关,在品种选择时应给予重视。

3.2 越橘果蒂痕和果实硬度性状遗传特点

本研究通过带柄采收模拟不形成果蒂痕,贮藏效果好,与常规脱柄贮藏形成鲜明对比,表明果实果蒂痕小对越橘果实贮藏意义重大,而目前生产上选择品种时重点考虑产量、品质等经济性状而忽略果蒂痕,因此,培育果蒂痕小而干的越橘品种,配合保鲜技术能有效提高越橘果实贮藏性。目前鲜见关于果蒂痕性状遗传规律的报道,遗传特点尚不明确。本研究利用正、反交群体对越橘果实果蒂痕遗传倾向进行分析,发现正交群体和反交群体后代果蒂痕大小平均值无明显差异,无父母本倾向,广义遗传力（H²）较高,表明果蒂痕大小的变异主要来自遗传效应,环境掩盖效应较小,遗传潜能大,可通过杂交选育小果蒂痕后代。

果实硬度是果实商品性和贮藏性的重要衡量指标之一^[36,37],对果实口感也有重要影响,就世界葡萄、草莓、越橘市场而言,消费者更青睐果肉脆硬/较硬的品种,因此果实硬度越来越受果树生产者和育种者的重视。以提高果实硬度为重要目标之一的草莓、葡萄、番茄等育种取得了重要进展,培育了很多果实硬度大、耐储运的新品种,对产业发展发挥了重要作用。浆果类果树中,在草莓上开展了果实硬度遗传的相关研究,但观点不尽一致。SPANGLO等^[38]认为果实硬度的遗传力较低,具显著的非加性效应;BARRITT^[39]则认为果实硬度具有较高的遗传力和较大的加性效应;李英慧等^[40]、祝朋芳等^[41]分别利用8个和10个杂交群体对草莓果实硬度进行遗传分析,果实硬度平均广义遗传力均为0.57,认为果实硬度属于多基因控制的数量性状,普遍存在正向非加性效应,草莓果实硬度较易通过杂交育种加以改善。番茄^[42]、梨^[28,43]中果肉硬度也存在相似遗传规律。本研究利用正、反交群体对越橘果实硬度的遗传倾向进行分析,结果显示正、反交后代果实硬度平均值无差异,均趋近低值亲本,受低值亲本影响,存在广泛的趋向硬度降低的劣质遗传趋势,此外,正、反交群体果实硬度的广义遗传力H²相对较低,说明该性状遗传效应小,易受环境影响,遗传中存在负向非加性效应。综上,越橘果实硬度存在广泛的趋低劣质遗传趋势,应避免选择果实硬度低的亲本,高硬度果实选择难度大,可能需要多代杂交聚合控制果实硬度的基因,从而获得果实硬度大的品种。

4 结论

果蒂痕大小和果实硬度是影响越橘果实贮藏的主要性状,培育果蒂痕小、果实硬度大的越橘品种对提高越橘果实贮藏性具有重要意义。果蒂痕大小和果实硬度表现为多基因控制的数量性状,其中果蒂痕大小存在趋中变异,无父母本倾向,变异主要来自遗传效应,遗传潜能大,超高亲遗传不明显;果实硬度平均值均趋近于低值亲本,存在遗传广泛的趋向硬度降低的劣质遗传趋势,且遗传效应小,易受环境影响。

致谢：

感谢北京科技大学刘冬成研究员对英文摘要的校正,感谢辽宁省果树科学研究所刘硕博士在分析软件使用中的指导。

参考文献原文顺序
文献年度倒序
文中引用次数倒序
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[本文引用: 2]

[7]

吉宁, 龙晓波, 李江阔, 曹森, 张鹏, 马超, 马立志, 王瑞. 1-MCP 结合臭氧处理对蓝莓低温保鲜效果的影响
食品工业科技, 2019(11): 302-307.

[本文引用: 2]

, LONG

X B

, LI

J K

, CAO

, ZHANG

, MA

L Z

, WANG

. Effect of 1-MCP coupling with ozone treatment on storage of blueberry at low temperature
Science and Technology of Food Industry, 2019(11): 302-307. (in Chinese)

[本文引用: 2]

[8]

巴良杰, 罗冬兰, 曹森, 尤勇刚, 余碧青, 王瑞. 1-MP结合⁶⁰Co γ辐照对蓝莓贮藏品质的影响
北方园艺, 2019(8): 118-124.

[本文引用: 2]

L J

, LUO

D L

, CAO

, YOU

Y G

, YU

B Q

, WANG

. Effects of 1-MCP combined with ⁶⁰Co γ irradiation on the quality of blueberry during storage
Northern Horticulture, 2019(8): 118-124. (in Chinese)

[本文引用: 2]

[9]

纪淑娟, 周倩, 马超, 程顺昌. 1-MCP处理对蓝莓常温货架品质变化的影响
食品科学, 2013, 35(2): 322-327.

[本文引用: 2]

S J

, ZHOU

, MA

, CHENG

S C

. Effect of 1-MCP treatment on quality changes of blueberry at room temperature
Food Science, 2013, 35(2): 322-327. (in Chinese)

[本文引用: 2]

[10]

CHIABRANDO

, GIACALONE

. Shelf life extension of highbush blueberry using 1-methylcyclopropene stored under air and controlled atmosphere
Food Chemistry, 2011, 126(4): 1812-1816.

DOI:10.1016/j.foodchem.2010.12.032 URL [本文引用: 2]

The potential of 1-methylcyclopropene for controlling ripening in 'Lateblue' blueberry fruit was explored. After harvest, blueberry fruits were exposed to 1-MCP (0.3 and 0.6 mu l l(-1)). After treatment, samples were stored in air at 0 degrees C for 35 days and in a controlled atmosphere (3 kPa O(2) + 11 kPa CO(2)) for 60 days. Quality parameters were monitored (weight loss, total soluble solids content, titratable acidity, firmness, anthocyanin content, phenolic content, total antioxidant capacity). Blueberries treated with 1-MCP showed a reduced weight loss during storage and a lower total soluble solid content compared to untreated fruit. High titratable acidity values were observed after controlled atmosphere storage, but no significant effect of 1-MCP on this parameter was observed. 1-MCP had no significant effects on anthocyanins, phenolics or antioxidant activities. (C) 2010 Elsevier Ltd.

[11]

郭丹, 韩英群, 郝义. 箱式气调保鲜对蓝莓果实贮藏期及货架期品质生理影响
西北农业学报, 2016, 25(12): 1829-1836.

[本文引用: 2]

GUO

, HAN

Y Q

, HAO

. Effect of box modified atmosphere storage on quality and physiology of blueberry during storage and shelf-life
Acta Agriculturae Boreali-occidentalis Sinica, 2016, 25(12): 1829-1836. (in Chinese)

[本文引用: 2]

[12]

刘萌, 范新光, 王美兰, 周志才, 张长峰, 张玉华. 不同包装方法对蓝莓采后生理及贮藏效果的影响
食品科学, 2013, 34(14): 346-350.

DOI:10.7506/spkx1002-6630-201314072 URL [本文引用: 2]

The present study investigated the influence of 1-methylcyclopropene (1-MCP) on fruit quality and reactive
oxygen species of blueberry fruit by multivariate analysis. The results showed that 1-MCP reduced the decay index, and
inhibited the increase of respiratory intensity as well as pH, glutathione (GSH) and malonaldehyde (MDA) of blueberry
fruit during storage at 1 ℃ and subsequent shelf life. The decline of H2O2 content was promoted at the earlier stage, but
effective delay was observed in fruits treated with 15 μL/L 1-MCP. Principal component analysis (PCA) showed that at the
earlier stage of storage, treatment with 5 μL/L 1-MCP significantly influenced the activities of polyphenol oxidase (PPO) and
peroxidase (POD) and the content of MDA, and affected GSH content, catalase (CAT) activity, pH and respiratory intensity during the
earlier period of shelf life. In contrast, POD activity and MDA content of fruit were mainly influenced by 15 μL/L 1-MCP at the later
stage of storage, and 5 μL/L1-MCP showed a significant impact on decay index and the activities of ascorbic peroxidase
(APX), lipoxygenase (LOX) and superoxide dismutase (SOD) of blueberry fruit during the later period of shelf life. Partial
least squares regression (PLSR) and path analysis showed that APX, LOX and SOD were the key factors to evaluate decay
index of blueberry fruit. The influences of respiratory intensity, the content of MDA and protein on decay index were
impacted by CAT activity and GSH content. In addition, the content of H2O2 was mainly influenced by the activities of SOD
and CAT, and the impact of MDA on H2O2 was manifested through GSH.

LIU

, FAN

X G

, WANG

M L

, ZHOU

Z C

, ZHANG

C F

, ZHANG

Y H

. Influence of different packaging methods on physiological properties of blueberry during postharvest storage
Food Science, 2013, 34(14): 345-350. (in Chinese)

DOI:10.7506/spkx1002-6630-201314072 URL [本文引用: 2]

孟宪军, 姜爱丽, 胡文忠, 田密霞, 刘程惠. 箱式气调贮藏对采后蓝莓生理生化变化的影响
食品工业科技, 2011, 32(9): 379-383.

[本文引用: 2]

MENG

X J

, JIANG

A L

, HU

W Z

, TIAN

M X

, LIU

C H

. Effects of plastic box modified atmosphere storage on the physiological and biochemical changes of postharvest blueberry fruits
Science and Technology of Food Industry, 2010, 32(9): 379-383. (in Chinese)

[本文引用: 2]

[14]

邹小波, 杨志坤, 石吉勇, 黄晓玮, 张文, Haroon

Elrasheid TAHIR

. 阿拉伯胶/白色玫瑰茄提取物复合涂膜对低温贮藏蓝莓保鲜效果的影响
食品科学, 2019, 40(7): 204-211.

[本文引用: 1]

ZOU

X B

, YANG

Z K

, SHI

J Y

, HUANG

X W

, ZHANG

, HAROON

E T

. Preservation effect of gum arabic edible coating incorporated with white roselle extract (
Hibiscus sabdariffa L.) on cold-stored blueberries. Food Science, 2019, 40(7): 204-211. (in Chinese)

[本文引用: 1]

[15]

李翠萍, 袁磊, 杨延存, 王茜, 周凯乐, 刘双, 李鹏程. 山梨醇提高壳聚糖季铵盐涂膜对蓝莓的保鲜效果
现代食品科技, 2018, 34(12): 117-121.

[本文引用: 1]

C P

, YUAN

, YANG

Y C

, WANG

, ZHOU

K L

, LIU

, LI

P C

. Effect of plasticizers on the preservation of blueberries by quaternary ammonium salt of chitosan coating
Modern Food Science and Technology, 2018, 34(12): 117-121. (in Chinese)

[本文引用: 1]

[16]

张群利, 崔琳琳, 郑权, 邬泽凯, 丁云闪, 覃懿琴, 张佳俐, 何美杉, 陈钧浦. 纤维素复合抗菌保鲜膜包覆对采后蓝莓的保鲜效果
食品工业科技, 2019, 40(19): 261-265.

[本文引用: 1]

ZHANG

Q L

, CUI

L L

, ZHENG

, WU

Z K

, DING

Y S

, TAN

Y Q

, ZHANG

J L

, HE

M S

, CHEN

J P

. Fresh-keeping effect of cellulose composite antibacterial preservative film on postharvest blueberry
Science and Technology of Food Industry, 2019, 40(19): 261-265. (in Chinese)

[本文引用: 1]

[17]

孙莎, 郜海燕, 熊涛, 陈杭君, 刘瑞玲, 吴伟杰. 五倍子提取液对蓝莓采后病害和品质的影响
林业科学, 2018, 54(6): 53-62.

[本文引用: 1]

SUN

, GAO

H Y

, XIONG

, CHEN

H J

, LIU

R L

, WU

W J

. The effect of galla chinensis extract on postharvest disease and storage quality of blueberry
Scientia Sivae Sinicae, 2018, 54(6): 53-62. (in Chinese)

[本文引用: 1]

[18]

LIN

S H

, WANG

, ZHANG

J H

, YANG

X L

. Initial screening of 10 plant essential oils in vitro antibacterial activity against blueberry pathogens
Agricultural Science & Technology, 2019, 20(3): 17-21.

[本文引用: 1]

[19]

王和涛. 丁香精油与壳聚糖涂膜对蓝莓的保鲜效果
食品工业, 2019, 40(1): 203-206.

[本文引用: 1]

WANG

H T

. Effect of clove essential oil and chitosan compound film coating on blueberry preservation
Food Industry, 2019, 40(1): 203-206. (in Chinese)

[本文引用: 1]

[20]

王磊明, 李洋, 张茜, 冯刚. 壳聚糖-肉桂精油复合膜对蓝莓保鲜效果的影响
食品科技, 2017, 42(9): 14-20.

[本文引用: 1]

WANG

L M

, LI

, ZHANG

, FENG

. Effects of composite membrane of chitosan and cinnamon essential oil on preservation of blueberries
Food Science and Technology, 2017, 42(9): 14-20. (in Chinese)

[本文引用: 1]

[21]

陶烨, 王琛, 高雅, 李晓平, 崔智博, 孟宪军. 不同剂量⁶⁰Co-γ辐照对蓝莓果实贮藏品质的影响
食品安全质量检测学报, 2017, 8(7): 2779-2785.

[本文引用: 1]

TAO

, WANG

, GAO

, LI

X P

, CUI

Z B

, MENG

X J

. Effects of different doses of ⁶⁰Co-γirradiation on storage quality of blueberry fruits
Journal of Food Safety and Quality, 2017, 8(7): 2779-2785. (in Chinese)

[本文引用: 1]

[22]

龙明秀, 吴凤玉, 田竹希, 刘敏, 何扬波, 李咏富, 石彬, 梁倩. ⁶⁰Co-γ辐照处理对蓝莓保鲜效果的影响
核农学报, 2019, 33(11): 2165-2176.

[本文引用: 1]

LONG

M X

, WU

F Y

, TIAN

Z X

, LIU

, HE

Y B

, LI

Y F

, SHI

, LIANG

. Effect of ⁶⁰Coγirradiation treatment on preservation effects of blueberry
Journal of Nuclear Agricultural Sciences, 2019, 33(11): 2165-2176. (in Chinese)

[本文引用: 1]

[23]

姬亚茹, 周福慧, 姜爱丽, 顾思彤, 胡文忠. 乙醇熏蒸处理对采后蓝莓果实品质的影响
包装工程, 2018, 13: 85-92.

[本文引用: 1]

Y R

, ZHOU

F H

, JIANG

A L

, GU

S T

, HU

W Z

. Effects of ethanol fumigation treatments on the quality of postharvest blueberry fruits
Packaging Engineering, 2018, 13: 86-92. (in Chinese)

[本文引用: 1]

[24]

廖嘉, 胡文忠, 权春善, 姬亚茹, 杨香艳, 龙娅, 李元政. 采后浆果熏蒸保鲜技术的研究进展
食品与发酵工程, 2019, 45(14): 277-284.

[本文引用: 1]

LIAO

, HU

W Z

, QUAN

C S

, JI

Y R

, YANG

X Y

, LONG

, LI

Y Z

. Fumigation preservation technology for postharvest berries: A review
Food and Fermentation Industries, 2019, 45(14): 277-284. (in Chinese)

[本文引用: 1]

[25]

刘有春, 陶承光, 魏永祥, 刘成, 王兴东, 刘威生, 杨艳敏. 越橘果实糖酸含量和不同发育阶段的变化及其与叶片中可溶性糖含量的相关关系
中国农业科学, 2013, 46(19): 4110-4118.

DOI:10.3864/j.issn.0578-1752.2013.19.017 URL [本文引用: 1]

【Objective】An experiment was conducted to study the characters of sugar and acid content and analyze the relationships of sugar content in fruits and leaves during fruit development, thus providing a theoretical basis for metabolic mechanism of sugar and acid and quality regulation, in blueberry.【Method】Five cultivars of three taxonomic types , including high bush blueberry, e.g. ‘Spartan’ and ‘Jersey’, half-high bush blueberry, e.g. ‘North country’ and ‘North blue’ and low bush blueberry, e.g. ‘Blomidon’ were used to investigate the seasonal variation of sugar and acid content in fruits and leaves at different fruit development stages by high performance liquid chromatography (HPLC). 【Result】The results indicated that the average content of total sugar was 102.04 mg•g-1 FW, the highest and lowest of total sugar content existed in ‘Spartan’ and ‘North country’, respectively. Glucose and fructose content increased during fruit maturation and the contents of them accounted for 97.90%-99.47% of total sugar content in mature fruit, the ratio of them was 1:1, whereas the sucrose and sorbitol contents decreased constantly from a low level during fruit maturation into traces of them in matured fruit. The average content of total acid was 7.10 mg•g-1 FW, in tested high bush blueberry and half-high bush, citric acid was the major organic acid, accounted for 76.94% of total acid in mature fruit which increased at early stage, and decreased quickly till fruit maturity, quinic acid and malic acid contents decreased gradually. There was a similar change trend and component in both low bush cultivars ‘Blomidon’ and half-high bush cultivar ‘North country’, with quinic acid being the main acid and it presented a decline tendency during the fruit development stage, the content was higher than other cultivars in matured fruit, citric acid and tartaric were the minor components; Sorbitol accounted for 67.28% of total sugar in leaves, and the content reached top at 42 d after full bloom (around 15 d before harvest) , then decreased quickly.【Conclusion】The component characters of organic acid in ‘Blomidon’ and ‘North country’ were obviously distinguished from high bush and half-high bush blueberry. Sorbitol was the main soluble sugars in leaves. The stage of around 15 days before harvest was the critical period of fruit expansion growth and sugar accumulation.

LIU

Y C

, TAO

C G

, WEI

Y X

, LIU

, WANG

X D

, LIU

W S

, YANG

Y M

. Fruit sugar and acid content, variation at different fruit development stages and their relationship with leaf soluble sugar content of blueberry
Scientia Agricultura Sinica, 2013, 46(19): 4110-4118. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2013.19.017 URL [本文引用: 1]

刘有春, 魏永祥, 王兴东, 刘成, 蒋明三, 张舵, 袁兴福, 陶承光. 南高丛越橘品种‘Sapphire’和北高丛品种‘Berkeley’正反交后代果实糖酸组分含量的遗传倾向
中国农业科学, 2014, 47(24): 4878-4885.

DOI:10.3864/j.issn.0578-1752.2014.24.011 URL [本文引用: 1]

【Objective】 The objective of present study is to analyze the effects of parents on inheritance of sugar and acid contents in blueberry fruit, to provide a basis for selecting parents in cross breeding. 【Method】 Sugar and acid contents of southern high bush blueberry ‘Sapphire’ ( high sugar and high acid type), northern high bush blueberry ‘Berkeley’ (middle sugar and low acid type) and their reciprocal cross progenies were investigated with high performance liquid chromatography (HPLC) method. 【Result】 Fructose and glucose were the major components of sugar, while citric acid and tartaric acid were the major components of acid. The reciprocal populations exhibited a wide phenotypic variation in fructose, glucose, total sugar, tartaric acid, citric acid and total acid contents where the coefficient of variation (CV) of sugar and acid contents were more than 15% and 30%, respectively, and the range of acid contents was higher than that of sugar contents, indicating that there was a bigger selecting potential for acid content. Mean sugar and acid contents of reciprocal cross progenies were lower than mid-parental values, showing a tendency of depression. The mean sugar contents of progenies inclined to low sugar parent, while the mean acid contents inclined to paternal parent value. The individual and total sugar contents presented a normal distribution, implying sugar contents are quantitative traits and controlled by polygenenes, while individual and total acid presented a skewed distribution, implying acid contents are quantitative traits and may be controlled by both major gene and polygenes. The broad sense heritability (H²) of fructose, glucose, total sugar, citric acid and total acid contents were ranged from 0.66 to 0.86, in which the highest was fructose (0.84 and 0.86 in the reciprocal populations, respectively), indicating that the variations of these traits mainly resulted from inheritance, and they were mainly affected by additive effect, while the H² of tartaric acid was the lowest (0.22 and 0.30 in the reciprocal populations, respectively), indicating that the variation of tartaric acid is greatly affected by environment factors. 【Conclusion】 The blueberry reciprocal populations exhibited a wide phenotypic variation in both sugar and acid contents, mean sugar and acid contents of progenies were lower than mid-parental values. The sugars contents presented a normal distribution, and were the quantitative traits and controlled by polygenenes. The mean sugar contents of progenies come close to low sugar parent, there exists maternal inheritance on the sugar contents, while the mean acid contents come close to paternal value and were mainly affected by paternal parent.

LIU

Y C

, WEI

Y X

, WANG

X D

, LIU

, JIANG

M S

, ZHANG

, YUAN

X F

, TAO

C G

. Inheritance tendency of sugar and acid contents in the reciprocal cross progenies’ fruits of Southern× Northern high Bush blueberry (Vaccinium)
Scientia Agricultura Sinica, 2014, 47(24): 4878-4885. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2014.24.011 URL [本文引用: 1]

CHEN

X S

, WU

, CHEN

M X

, HE

T M

, FENG

J R

, LIANG

, LIU

, YANG

H H

, ZHANG

L J

. Inheritance and correlation of self-compatibility and other yield components in the apricot hybrid F1 populations
Euphytica, 2006, 150(12): 69-74.

DOI:10.1007/s10681-006-9094-7 URL [本文引用: 1]

[28]

崔艳波, 陈慧, 乐文全, 张树军, 伍涛, 陶书田, 张绍玲. ‘京白梨’与‘鸭梨’正反交后代果实性状遗传倾向研究
园艺学报, 2011, 38(2): 215-224.

URL [本文引用: 2]

‘Jingbaili’，‘Yali’ pear cultivars and their F1 offspring of reciprocal crossing were used to study genetic tendency of some fruit characters. The results showed that，most of the fruit characters were quantitative and controlled by polygenic. Fruit size was observed to be smaller in F1 generation，with the average transmitting ability and average coefficient of variation as 77.14% and 38.26% respectively. Compared with parent’s fruit，increase of soluble solids and soluble sugar content were obtained together in the hybrid fruits，97.25% and 35.28%，higher than those of parents respectively basing on average values. Firmness and titratable acid also tended to be higher than parents，with the average transmitting ability and coefficient of variation as 123.81% and 51.1% respectively. Same for content of stone cells，it was significant higher in F1 generation，and the value of average transmitting ability was up to 182.62%. As regards the vitamin C content，fruit of F1 generation had opposite genetic tendency when‘Jingbaili’or‘Yali’was as pollen parent.

CUI

Y B

, CHEN

, LE

W Q

, ZHANG

S J

, WU

, TAO

S T

, ZHANG

S L

. Studies on genetic tendency of fruit characters in reciprocal crosses generation between ‘Jingbaili’ and ‘Yali’ pear cultivars
Acta Horticulturae Sinica, 2011, 38(2): 215-224. (in Chinese)

URL [本文引用: 2]

刘有春, 鄂辉邦, 刘成, 王兴东, 杨艳敏, 孙斌, 张舵, 袁兴福, 魏永祥. 越橘半同胞系杂交后代果实糖酸性状的变异和遗传倾向
果树学报, 2016, 33(6): 664-675.

[本文引用: 1]

LIU

Y C

, E

H B

, LIU

, WANG

X D

, YANG

Y M

, SUN

, ZHANG

, YUAN

X F

, WEI

Y X

. Variability and inheritance of sugar and acid content in blueberry (Vaccinium) fruit of half-sib crosses
Journal of Fruit Science, 2016, 33(6): 664-675. (in Chinese)

[本文引用: 1]

[30]

WICKHAM

. ggplot2: elegant graphics for data analysis
Springer. 2016

[本文引用: 1]

[31]

周会玲, 李嘉瑞. 葡萄果实结构与耐贮性关系的研究
中国农学通报, 2005, 21(7): 239-241.

URL [本文引用: 1]

In this paper, the Relationship between attachment of stem to the berries, turgor pressure and fruit structure in four table grapes were studied of Kyoho, Red Globe,Autumn black and Autum Red grape.The result showed that attachment of stem to the berries was positvely correlated to the diameter of fruit stalk, diameter of the fruit stalk end and total fruit stalk binding area but negatively to the total berry weight per unit area of stalk and stalk binding area. Turgor pressure not only decided by flesh character but also by the berry size,but not related to the pericarp thickness. The fruit stalk xyloidization extent effect the storability of grape more than the stalk to stem ratio.,but the attachment of stem to the berries was not related to the turgor pressure.

ZHOU

H L

, LI

J R

. The relationship between fruit structure and storability in table grapes
Chinese Agricultural Science Bulletin, 2005, 21(7): 239-241. (in Chinese)

URL [本文引用: 1]

JEWEL

. Maintaining fruit firmness
[D]. Arizona: University of Arizona, 2004.

[本文引用: 1]

[33]

王金英, 马中国, 宗灿华. 果胶的提取与应用
中国林副特产, 2000, 5(2): 17-l8.

[本文引用: 1]

WANG

J Y

, MA

Z G

, ZONG

C H

. Extraction and application of pectin
Quarterly of Forest By-Product and Speciallty in China, 2000, 5(2): 17-18. (in Chinese)

[本文引用: 1]

[34]

魏宝东, 姜炳义, 冯辉. 番茄果实货架期硬度变化及其影响因素的研究
食品科学, 2005, 26(3): 249-252.

URL [本文引用: 2]

The fruit hardness degree of tomato and the factors affected were studied during the shelf phase with Baili, Katelina, R144, new tomatoes types all having hard fruit flesh and Gailun, TF997, TF998, new varieties breeded in Xingguang Seed Co.Ltd., as experimental materials, and with Lisheng Num. 8 as a check material. The results show that the fruit hardness degree and flesh of fruit hardness degree are the index to evaluate the lifespan of the fruit during shelf phase. The content of the insoluble pectin and the soluble pectin are the direct factors affecting both of them. Moreover, the fruit hardness degree and the flesh of fruit hardness degree have a positive correlation with the content of insoluble pectin as well as they have a negative correlation with the content of soluble pectin, both of them at a very significant level. PG and PE are the indirect factors affecting the softening of the fruit.

WEI

B D

, JIANG

B Y

, FENG

. Studies on changes of hardness degree of tomato fruit and factors affected shelf-Life
Food Science, 2005, 26(3): 249-252. (in Chinese)

URL [本文引用: 2]

张会芳, 刘桂珍, 张超. 草莓果实硬度影响因素及其调控措施研究进展
长江蔬菜, 2009(12): 4-7.

URL [本文引用: 1]

草莓采摘后极易软化腐烂,草莓的贮藏品质与果实硬度密切相关.概述了草莓果实软化机理,详述了草莓果实硬度的影响因素及调控措施.

ZHANG

H F

, LIU

G Z

, ZHANG

. Research progress on influencing factors and control measures of strawberry fruit firmness
Changjiang Vegetable, 2009(12): 4-7. (in Chinese)

URL [本文引用: 1]

草莓采摘后极易软化腐烂,草莓的贮藏品质与果实硬度密切相关.概述了草莓果实软化机理,详述了草莓果实硬度的影响因素及调控措施.

[36]

彭永彬, 李玉, 徐鹏程, 王晨, 上官凌飞. 葡萄果实硬度及影响硬度的主要因素
浙江农业学报, 2014, 26(5): 1227-1234.

URL [本文引用: 2]

The firmness of table grapes is one of the main factors that determine the eating quality of grapes， which is subjected to the increasing attention of the fruit producers and breeders. Many studies have been carried out on berry firmness. For a better understanding of the grape firmness， this paper summarized the fruit firmness variation and influencing factors to promote the further study of grape fruit firmness. Firstly， the fruit firmness and measuring method were introduced. Secondly， we expounded the changes of fruit firmness in different growth periods. Various influencing factors were summarized in the end. Internal factors included fruit cell structure and special ions， and exogenous hormones and external conditions belonged to external factors.

PENG

Y B

, LI

, XU

P C

, WANG

, SHANGGUAN

L F

. Grape fruit firmness and the main influencing factors
Acta Agriculturae Zhejiangensis, 2014, 26(5): 1227-1234. (in Chinese)

URL [本文引用: 2]

王力荣, 朱更瑞, 方伟超. 桃(Prunus persica L.)种质资源果实数量性状评价指标探讨
园艺学报, 2005, 32(1): 1-5.

URL [本文引用: 1]

We try to establish a new peach descrip tor system based on the statistics analysis, international Peach Descriptor from IBPGR in 1984, and Chinese peach descriptor in 1990. Fruit quantitative characters of 282 - 491 peach cultivars were investigated atNational Peach Genetic Collection in Zhengzhou
in China from 1986 - 2002. Those characters includes average fruit mass, stone mass, soluble solid content, soluble sugar content, soluble acid content, vitamin C, firmnesswith shin and firmnesswithout skin.Grade index and reference cultivarswere given by statistical data for peach descrip tors. These grade indexes were recorded on 1 - 5 scale. In generally, two reference cultivars for each grade were given, one USA cultivar, another Chinese cultivar. We try to use these as the reference cultivars, which are planted or used by
breeders widely in China.

WANG

L R

, ZHU

G R

, FANG

W C

. The evaluating criteria of some fruit quantitative characters of peach (Prunus persica L.) genetic resource
Acta Horticulturae Sinica, 2005, 32(1): 1-5. (in Chinese)

URL [本文引用: 1]

SPANGLO

L P S

, HSU

C S

, FEJER

S O

. Heritability and genetic variance components for 20 fruit and plant characters in the cultivated strawberry
Genome, 1971, 13: 443-456.

[本文引用: 1]

[39]

BARRITT

B H

. Breeding strawberry for fruit firmness
Journal of the American Society for Horticultural Science, 1979, 104(5): 663-665.

[本文引用: 1]

[40]

李英慧, 王火旭, 任秀云. 草莓果实硬度遗传和选择研究
辽宁师范大学学报:自然科学版, 2000, 20(3): 316-318.

[本文引用: 1]

Y H

, WANG

H X

, REN

X Y

. Study on heredity and selection of fruit firmness of strawberry
Journal of Liaoning Normal University (Natural Science Edition), 2000, 20(3): 316-318. (in Chinese)

[本文引用: 1]

[41]

祝朋芳, 陈长青. 草莓两个经济性状遗传特性的研究
北方果树, 2004(3): 8-9.

URL [本文引用: 1]

该文研究了草莓最大果重、果实硬度的遗传特性.结果表明,最大果重遗传存在着负向非加性效应,杂交后代最大果重呈退化趋势;果实硬度遗传普遍存在着正向非加性效应,杂交后代中易选出果实硬度大的株系.

ZHU

P F

, CHEN

C Q

. A study on genetic traits of two economic characters in strawberry
Northern Fruits, 2004(3): 8-9. (in Chinese)

URL [本文引用: 1]

该文研究了草莓最大果重、果实硬度的遗传特性.结果表明,最大果重遗传存在着负向非加性效应,杂交后代最大果重呈退化趋势;果实硬度遗传普遍存在着正向非加性效应,杂交后代中易选出果实硬度大的株系.

[42]

何艳龙. 番茄果实硬度遗传规律研究
北方园艺, 2016(14): 17-22.

[本文引用: 1]

Y L

. Study on genetic low of fruit firmness of tomato
Northern Horticulture, 2016(14): 17-22. (in Chinese)

[本文引用: 1]

[43]

赵亚楠, 欧春青, 王斐, 张艳杰, 马力, 姜淑苓. 梨F₁代群体果实性状的遗传分析及相关性研究
西北农业学报, 2018, 27(12): 1811-1818.

[本文引用: 1]

ZHAO

Y N

, OU

Q C

, WANG

, ZHANG

Y J

, MA

, JIANG

S L

. Genetic analysis and correlation study on pear fruit characteristics in F₁ generation
Acta Agriculturae Boreali-occidentalis Sinica, 2018, 27(12): 1811-1818. (in Chinese)

[本文引用: 1]