Genome-wide identification of the CONSTANS-LIKE(COL)family and mechanism of fruit senescence regulation by PpCOL8 in sand pear(Pyrus pyrifolia)

Pyrus pyrifolia Nakai‘Whangkeumbae'is a sand pear fruit with excellent nutritional quality and taste.However,the industrial development of pear fruit is significantly limited by its short shelf life.Salicylic acid(SA),a well-known phytohormone,can delay fruit senescence and improve shelf life.However,the mechanism by which SA regulates CONSTANS-LIKE genes(COLs)during fruit senescence and the role of COL genes in mediating fruit senescence in sand pear are poorly understood.In this study,22 COL genes were identified in sand pear,including four COLs(Pp COL8,Pp COL9a,Pp COL9b,and Pp COL14)identified via transcriptome analysis and 18 COLs through genome-wide analysis.These COL genes were divided into three subgroups according to the structural domains of the COL protein.Pp COL8,with two B-box motifs and one CCT domain,belonged to the first subgroup.In contrast,the other three Pp COLs,Pp COL9a,Pp COL9b,and Pp COL14,with similar conserved protein domains and gene structures,were assigned to the third subgroup.The four COLs showed different expression patterns in pear tissues and were preferentially expressed at the early stage of fruit development.Moreover,the expression of Pp COL8 was inhibited by exogenous SA treatment,while SA up-regulated the expression of Pp COL9a and Pp COL9b.Interestingly,Pp COL8 interacts with Pp MADS,a MADS-box protein preferentially expressed in fruit,and SA up-regulated its expression.While the production of ethylene and the content of malondialdehyde(MDA)were increased in Pp COL8-overexpression sand pear fruit,the antioxidant enzyme(POD and SOD)activity and the expression of Pp POD1 and Pp SOD1 in the sand pear fruits were down-regulated,which showed that Pp COL8 promoted sand pear fruit senescence.In contrast,the corresponding changes were the opposite in Pp MADS-overexpression sand pear fruits,suggesting that Pp MADS delayed sand pear fruit senescence.The co-transformation of Pp COL8 and Pp MADS also delayed sand pear fruit senescence.The results of this study revealed that Pp COL8 can play a key role in pear fruit senescence by interacting with Pp MADS through the SA signaling pathway.

Identification of Self-Incompatibility Genotypes in Some Sand Pears (Pyrus pyrifolia Nakai) by PCR-RFLP Analysis

The identification of self-incompatibility genotype （S-genotype） will be useful for selection of pollinizers and design of crossing in cultivar improvement of sand pear. This paper reported the identification of self-incompatibility genotypes of seven Chinese and two Japanese sand pear cultivars using PCR-RFLP analysis and S-RNase sequencing. The Sgenotypes of these cultivars were determined as follows： Huali 1 S1S3, Shounan S1S3, Xizilti S1S4, Qingxiang S3S7, Sanhua S2S7, Huangmi （Imamuranatsu） S1S6, Huali 2 S3S4, Baozhuli S7S33, Cangxixueli S5S15. S-RNase alleles （S1 to S9） in sand pear could be identified effectively by PCR-RFLP analysis.

Isolation and Characterization of a Cinnamoyl CoA Reductase Gene(CCR) in Pear(Pyrus pyrifolia)

Pear is a popular and commercially important fresh fruit, and its texture is related to the presence of sclereid formatted by parenchyma cell with lignification in vascular plants. Previous studies have demonstrated that content of lignin may be regulated by cinnamoyl CoA reductase(CCR) in various plants. However, the function of CCR in pears remains very limited. In the present study, we isolated a cDNA encoding CCR(PpCCR, GenBank accession No. KF999958) and its promoter(proPpCCR) from Whangkeumbae pear to investigate the function of CCR in lignin biosynthesis. PpCCR-GFP expressed in rice mesophyll protoplast demonstrated that PpCCR-GFP was localized in the cytoplasm, indicating that CCR may function in cytoplasm without localization signals. In transgenic plants carrying PpCCR, we observed higher lignin content compared with that in wild type plants, further suggesting that PpCCR can affect the lignin contents through regulating lignin biosynthesis in Arabidopsis thaliana. More studies in other plants are needed to confirm our conclusion.

砂梨(Pyrus pyrifolia)过敏原蛋白基因(Ppmal)的克隆与表达分析

【目的】分离和克隆梨过敏原蛋白基因Ppmal(Gen Bank登录号为KP008110),探讨其在梨果实发育过程中的功能。【方法】以砂梨品种‘翠冠’[Pyrus pyrifolia(Burm.f.)Nakai]为试材,在盛花期后30 d对植株果柄进行涂抹赤霉素处理,利用同源克隆和RACE方法克隆目的基因全长序列,并通过实时荧光定量技术和半定量技术分析该基因在梨不同组织以及梨果实发育过程中的表达变化。【结果】Ppmal的全长是906 bp,含有480 bp的开放阅读框(ORF),编码159个氨基酸,预测的蛋白质相对分子质量为17.56 k D,等电点为5.62。生物信息学分析显示该基因没有信号肽序列,没有跨膜结构域,具亲水性。与其他物种的过敏原蛋白基因核苷酸序列同源性超过75%,与苹果和西洋梨编码的氨基酸序列同源性分别高达97.48%和96.23%,进化树分析表明该基因与苹果的进化关系最近。同时,定量结果显示经过赤霉素处理后的果实中该基因的表达量随着果实的生长发育呈现先降低后升高的趋势,并且具有组织差异表达的特点,其表达量在叶片中最大,其次是嫩叶,再次是花,枝条最低。【结论】获得梨Ppmal基因全长,对该基因在砂梨品种‘翠冠’发育过程中的表达分析显示,Ppmal受到赤霉素的调控,并在砂梨果实膨大期发生上调表达。

Relationship Between Anthocyanin Biosynthesis and Related Enzymes Activity in Pyrus pyrifolia Mantianhong and Its Bud Sports Aoguan

The aim of this article is to study the relationship between biosynthesis of anthocyanin and activities of phenylalanine ammonia lyase （PAL）, chalcone ismoerase （CHI） enzymes in Pyrus pyrifolia. Changes in the level of anthocyanin and the activities of enzymes of anthocyanin biosynthesis including PAL, CHI were studied in the pericarp of Pyrus pyrifolia Aoguan and Mantianhong during the period of pigment formation. Bagging treatment was also carried out to manipulate the synthesis of anthocyanin and the activities of related enzymes during the period of pigment formation. The results demonstrated that the level of anthocyanin of Aoguan was higher than that of Mantianhong. However, the content of anthocyanins has the similar changing trend in Aoguan and Mantianhong, highest anthocyanin concentrations of two varieties appeared in immature fruit and faded toward harvest. Meanwhile, similar changing trends of activities of PAL and CHI were also observed in both varieties. Aoguan has a lower activity of PAL than Mantianhong, whereas activity of CHI in Aoguan was higher than that in Mantianhong. Activity of PAL decreased during the period of pigment formation and was apparently not limiting to color development, whereas CHI activity increased at the same period and was closely related to the synthesis of anthocyanin. The results of bagging treatment showed that bagging treatment inhibited the activity of CHI, as well as the synthesis of anthocyanin, whereas debagging enhanced both the activity of CHI and synthesis of anthocyanin. The activity of CHI in debagging Aoguan pericarp was higher than the untreated Aoguan. However, effect of bagging treatment toward PAL activity was not obvious. Anthocyanin of bagging treated Aoguan decreased toward harvest. The content of anthocyanin of Pyruspyrifolia increased at the beginning of fruit coloration period and decreased toward fruit harvest. Activity of PAL was apparently not limiting to color development, whereas CHI activity was closely related to the synthesis of anthocyanin. Debagging enhanced both the activity of CHI and anthocyanin synthesis. Bagging treatment also proved that degradation of anthocyanin was not induced only by light. Light appeared to have two opposing effects in pears： it is required for anthocyanin synthesis and also for apparently increasing red color loss through increased degradation of anthocyanin.

Genomic selection of eight fruit traits in pear

Genomic selection (GS) has the potential to improve selection efficiency and shorten the breeding cycle in fruit tree breeding. In this study,we evaluated the effect of prediction methods, marker density and the training population (TP) size on pear GS for improving its performance and reducing cost. We evaluated GS under two scenarios:(1) five-fold cross-validation in an interspecific pear family;(2) independent validation. Based on the cross-validation scheme, the prediction accuracy (PA) of eight fruit traits varied between 0.33 (fruit core vertical diameter)and 0.65 (stone cell content). Except for single fruit weight, a slightly better prediction accuracy (PA) was observed for the five parametrical methods compared with the two non-parametrical methods. In our TP of 310 individuals, 2 000 single nucleotide polymorphism (SNP) markers were sufficient to make reasonably accurate predictions. PAs for different traits increased by 18.21%-46.98%when the TP size increased from 50to 100, but the increment was smaller (-4.13%-33.91%) when the TP size increased from 200 to 250. For independent validation, the PAs ranged from 0.11 to 0.45 using rrBLUP method. In summary, our results showed that the TP size and SNP numbers had a greater impact on the PA than prediction methods. Furthermore, relatedness among the training and validation sets, and the complexity of traits should be considered when designing a TP to predict the test panel.

PbrARF4 contributes to calyx shedding of fruitlets in ‘Dangshan Suli’ pear by partly regulating the expression of abscission genes

Fruitlet calyx shedding in pear plants is apparently regulated via numerous pathways that involve both environmental triggers and phytohormones cues such as auxin. In this study, we found at 10 days after full bloom (DAFB) higher levels of indoleacetic acid (IAA) and tryptophan (Trp) in calyx persistence fruitlet (CPF) than calyx shedding fruitlet (CSF) ofDanshan Suli’ pear (Pyrus bretschneideri Rhed.). Consisting with this, the activity of indolealdehyde oxidase (IAAIdO), which promotes IAA synthesis, was remarkably increased, and that of peroxidase(POD), which degrades IAA, dropped markedly in CPF but not in CSF. Further, qRT-PCR results revealed that most of 31 PbrARFs (encoding auxin response factors) in Pyrus bretschneideri were highly expressed in CPF, whereas PbrARF4, PbrARF24 and PbrARF26 were significantly downregulated in CPF vis-a-vis CSF. Phylogenetic analysis revealed that 6 PbrARFs clustered in the group III, where PbrARF4 showed the closest affinity with AtARF1 that promotes organ abscission, indicating a putative role of PbrARF4 in mediating the process of calyx shedding in pear. In fact, the ectopic overexpression of PbrARF4 in Solanum lycopersicum resulted in an earlier-formed and deeper abscission layer (AL) in the transgenic plants, whose calyxes were more prone to wilt at the mature red stage (MR) compared with the control plants (wild-type). More importantly, expression levels of the abscission genes SILS and Sl Cel2 in transgenic plants overexpressing PbrARF4 were significantly upregulated in comparation with the WT, whereas those of Sl BI and Sl TAPG2 were considerably inhibited. Further, PbrJOINTLESS and PbrIDA,the two genes related to calyx shedding in pear, were up-regulated more in CSF than CPF. The findings contribute to a better understanding of PbrARFs involved in fruitlet calyx shedding of pear, which could prove beneficial to improving the quality of pear fruit.

苹果梨（Pyrus Pyriforia （Burm） Nakai，cv．Pingguoli）果形畸变原因的探讨

本文研究了苹果梨果实畸变的原因，结果表明：果形偏斜与座果过多在生长期中互相挤压有关。果形严重畸变则是授粉不足或种子发育与生长不良的结果。这种变化在冬果梨的研究中也获得同样结果。对座果过多的果树可用疏花疏果的方法减少座果率。

Involvement of long non-coding RNAs in pear fruit senescence under high-and low-temperature conditions

Pear fruit senescence under high-and low-temperature conditions has been reported to be mediated by microRNAs.Long non-coding RNAs(lncRNAs),which can function as competing endogenous RNAs that interact with microRNAs,may also be involved in temperature-affected fruit senescence.Based on the transcriptome and microRNA sequencings,in this study,3330 lncRNAs were isolated from Pyrus pyrifolia fruit.Of these lncRNAs,2060 and 537 were responsive to high-and low-temperature conditions,respectively.Of these differentially expressed lncRNAs,82 and 24 correlated to the mRNAs involved in fruit senescence under high-and low-temperature conditions,respectively.Moreover,three lncRNAs were predicted to be competing endogenous RNAs(ceRNAs)that interact with the microRNAs involved in fruit senescence,while one and two ceRNAs were involved in fruit senescence under high-and low-temperature conditions,respectively.A dual-luciferase assay showed that the interaction of an lncRNA with a microRNA disrupts the action of the microRNA on the expression of its target mRNA(s).Furthermore,four alternative splicing-derived lncRNAs interacted with miR172i homologies(Novel_88 and Novel_69)to relieve the repressed expression of their target and produce an miR172i precursor.Correlation analysis of microRNA expression suggested that Novel_69 is likely involved in the cleavage of the pre-miR172i hairpin to generate mature miR172i.Taken together,lncRNAs are involved in pear fruit senescence under high-or low-temperature conditions through ceRNAs and the production of microRNA.

Genome-wide identification of the mitogen-activated protein kinase kinases in pear and their functional analysis in response to black spot

The mitogen-activated protein kinase(MAPK)cascade is crucial to plant growth,development,and stress responses.MAPK kinases(MAPKK)play a vital role in linking upstream MAPKK kinases(MAPKKK)with the downstream MAPK.Black spot is one of the most serious fungal diseases of pear which is an important part of the fruit industry in China.The MAPKK genes have been identified in many plants,however,none has been reported in pear(Pyrus bretschneideri).In order to explore whether MAPK gene of pear is related to black spot disease,we designed this experiment.The present study investigated eight putative PbrMAPKK genes obtained from the Chinese white pear genome.The phylogenetic analysis revealed that PbrMAPKK genes were divided into A,B,C,and D groups.These PbrMAPKK genes are randomly distributed on 7 out of 17 chromosomes and mainly originated from the whole-genome duplication(WGD)event.The expression analysis of PbrMAPKK genes in seven pear tissues and the leaves of susceptible and resistant varieties after Alternaria alternata infection by quantitative real-time PCR(qRT-PCR)identified seven candidate genes associated with resistance.Furthermore,virus-induced gene silencing(VIGS)indicated that PbrMAPKK6 gene enhanced resistance to pear black spot disease in pear.

The HY5 transcription factor negatively regulates ethylene production by inhibiting ACS1 expression under blue light conditions in pear

Ethylene is the main factor controlling fruit ripening of pear(Pyrus ussuriensis).Ethylene production rate is negatively correlated with fruit shelf life;therefore,it is important to decrease the ethylene levels for optimal fruit storage.Here,we observed that blue light treatment could inhibit ethylene production and promote the expression of ELONGATED HYPOCOTYL 5(PuHY5),a basic leucine zipper domain(bZIP)transcription factor.The following studies showed that PuHY5 could bind to the promoter of ACC synthase 1(PuACS1),a rate-limiting enzyme in ethylene biosynthesis,and inhibit its expression.For pears in which Pu HY5 was silenced,the ethylene production and PuACS1 expression were much higher than those in the control fruit.These results demonstrated that blue light inhibited ethylene production through the induction of Pu HY5 in pear.Our finding provides a new method for prolonging fruit shelf life.

两个秋子梨果实品质分析

为探究两个秋子梨品种差异,更好地对其进行开发利用,以红丰和早黄成熟期果实为试材,利用显著性分析、相关性分析、主成分分析和聚类分析等方法对两个不同品种秋子梨果实品质进行分析。结果表明,两种梨的果实在果实外观、果肉风味、果实香气、果汁含量上有所不同,在外观品质上红丰梨单果重为212.96 g、横径为75.49 mm、纵径为70.25 mm、果梗长度为36.48 mm,这4个指标分别达到了早黄梨的171.13%、119.33%、121.29%和113.19%;红丰梨的果实硬度显著低于早黄梨,为早黄梨的79.37%。因此红丰梨相较于早黄梨果形较大、果实较重、果肉较软,适合喜好大果、软果的消费者选择;在营养成分上红丰梨的可溶性固形物、可溶性糖、矿质元素含量均显著高于早黄梨,其中可溶性固形物含量为11.10%、矿质元素含量为3748.49 mg·kg^(-1),而可溶性糖含量为124.61 mg·g^(-1),达到了早黄梨可溶性糖含量的206.89%;果实的单果重、果实横径、果实纵径、果实硬度4项形态指标与可溶性糖、可溶性固形物、维生素C、矿质元素呈现出显著相关性,与可滴定酸呈极显著相关;在矿质元素含量上,红丰梨果肉和果皮中矿质元素Mg、P、K、Ca、Fe的含量以及12种矿质元素的总含量均高于早黄梨。对两个品种果实品质综合比较得出,红丰梨果实品质优于早黄梨,更适合在生产上开发利用。

Strategies for timing nitrogen fertilization of pear trees based on the distribution, storage, and remobilization of ^(15)N from seasonal application of (^(15)NH_4)_2SO_4

In order to improve the management of nitrogen(N) fertilization in pear orchards, we investigated the effects of application timing on the distribution, storage, and remobilization of N in mature pear trees in a field experiment at Jingtai County, Gansu Province, China. Nine trees were selected for the experiment and each received equal aliquots of 83.33 g N in the autumn, spring, and summer, with ^(15)N-labeled(NH_4)_2SO_4 used in one of the aliquots each season. Results showed that the(^(15)NH_4)_2SO_4 applied in the autumn remained in the soil during the winter. In the following spring this N was absorbed and rapidly remobilized into each organ, especially new organs(leaves, fruit and new shoots). The ^(15)N supplied in spring was rapidly transported to developing fruit between the young fruit and fruit enlargement stages. ^(15)N from the summer application of fertilizer was mainly stored in the coarse roots over the winter, then was mobilized to support growth of new organs in spring. In conclusion, for pear trees we recommend that the autumn application of N-fertilizer be soon after fruit harvest in order to increase N stores in fine roots. Spring application should be between full bloom and the young fruit stages to meet the high N demands of developing fruit. Summer application of fertilizer at the fruit enlargement stage does not contemporaneously affect the growth of pears, but increases the N stored in coarse roots, and in turn the amount available for remobilization in spring.

Genome-wide identification and co-expression analysis of GDSL genes related to suberin formation during fruit russeting in pear

Glycine-aspartic acid–serine-leucine(GDSL)type lipases/esterases genes play critical roles in plant development and are related to the responses to abiotic and biotic stress.However,little is known about the GDSL family in pear(Pyrus spp.).Studies have shown GDSL-domain proteins play key roles in suberin deposition.Suberin deposition in the fruit epidermis,also called russeting,is an important defect that negatively affects consumer's appeal in some fruit species,such as pear,apple and grapevine.Fruit russeting is mainly associated with cuticle microcracking and suberin accumulation in the inner part of the epidermal cell walls.To gain insight into the role of the GDSL gene family in suberin deposition and russet development in pear,we performed a genome-wide characterization of the GDSL family,including their identification,chromosomal localization,phylogenetic relationships,and expression patterns,in different tissues/organs in pear.One hundred and thirteen GDSL-type lipases/esterases genes were identified in the pear genome,and a phylogenetic analysis revealed that GDSL family can be classified into four distinct groups.Thirty GDSL genes were co-expressed with five homolog pear genes of three well-known suberin biosynthesis Arabidopsis genes(AtGPAT5,AtASFT,and AtCYP86B1)in the transcriptional co-expression network during pear fruit development.Among the 30 co-expressed GDSL genes,twelve genes were further analyzed by quantitative Real-time PCR,and the results showed the expression levels of the 12 genes were different between the russet exocarp and green exocarp of sand pear at different fruit development stages.Our study provides a detailed overview of the GDSL gene family and lays the foundation for future functional characterization of GDSL genes in P.bretschneideri.

砂梨品种果实冷藏期间质地及品质差异性评价

目的:确定砂梨品质形成及劣变关键点。方法:以早生新水、沪晶梨18、沪晶梨67为试材,将成熟度一致的3个品种的梨果实于(1.0±0.5)℃、相对湿度85%~90%的冷库中贮藏60 d,每10 d测定果实品质和10个电子鼻传感器的感应值。结果:沪晶梨18和早生新水为典型的果糖积累型,沪晶梨67为山梨醇积累型,3个品种梨均为苹果酸积累型。冷藏期间,沪晶梨67可保持较高的果实带皮硬度、果肉组织硬度、脆性和咀嚼性;含有较高的葡萄糖、山梨醇和总糖含量,可滴定酸、苹果酸含量较稳定;贮藏风味浓郁,受冷藏时间调控小,耐贮性佳。沪晶梨18的果皮特性、紧实度和脆性优于早生新水,拥有较高的果糖、苹果酸含量和果糖/总糖,可保持较高的甜酸度,贮藏风味浓郁。早生新水易软化,可滴定酸含量呈急剧下降趋势,综合风味受贮藏时间调控大,不耐贮。早生新水和沪晶梨67梨果实挥发性化合物发生了明显变化,特别是氮氧化合物、甲烷、硫化物、乙醇等挥发性化合物。结论:早生新水、沪晶梨18、沪晶梨67的安全冷藏期分别为30,40,60 d。

梨火疫病菌在库尔勒香梨枝条中的侵染定殖和扩展特征研究

【目的】示踪梨火疫病菌Erwinia amylovora在库尔勒香梨(Pyrus sinkiangensis Yu)(简称香梨)枝条组织中侵染定殖和扩展特征,为梨火疫病的有效防治提供依据。【方法】以分离获得梨火疫病强致病力菌株E.a001为材料,采用热激法将携带有绿色荧光蛋白基因(green fluorescent protein,GFP)质粒phc60-gfp转化E.a001,获得了能发出强烈绿色荧光的标记菌株E.a001-gfp。利用该标记菌株接种香梨枝条,检测、观察其在香梨枝条组织中的侵染定殖和扩展动态。【结果】针刺接种和喷雾接种E.a001-gfp引起枝条发病的最低浓度分别为106cfu·mL^(-1)和107cfu·mL^(-1)。相同浓度的病原菌接种,喷雾接种较针刺接种后发病显症时间延迟2~3 d。梨火疫病菌侵染枝条产生的坏死病斑的扩展速率随病原菌浓度提高而升高;病菌在1年生枝条上的生长扩展速度显著快于在2年生枝条内的扩展速度。梨火疫病菌入侵后主要存在于枝条维管系统中,大量分布于皮层薄壁细胞和韧皮部薄壁细胞间隙,少量存在于木质部导管中。在香梨发病枝条上,病健交界处的活菌数量最高,可达到108cfu·g^(-1),未显症部位0~10 cm处病原菌为105cfu·g^(-1),但在香梨枝条病斑外未显症部位10~15 cm处均未分离出病原菌。【结论】研究明确了梨火疫病菌在香梨枝条组织中的扩展距离、迁移时间及速率,以及分布及定殖量,为病枝的准确修剪和制定病害的综合防控措施提供了科学依据。

早熟砂梨苏翠1号在湖北省钟祥市的引种与栽培表现

早熟砂梨[Pyrus pyrifolia(Burm.F.)Nakai]新品种苏翠1号于2019年引入湖北省钟祥市梨产区进行引种栽培。结果表明,该品种在钟祥市表现为早果性好,成熟期早,果个大,石细胞少,风味甘甜,品质优,丰产、稳产,综合性状优良,是适合钟祥市梨产区发展的极早熟优良砂梨品种。

'玉露香’与’新世纪’杂交后代果实性状遗传研究

以’玉露香’与’新世纪’及94株杂交后代F1为材料,研究了8个果实重要性状的遗传倾向。结果表明:杂交后代平均单果重小于亲中值,遗传传递力为79.47%;果实硬度、可溶性固形物和可溶性糖大于亲中值,呈增大趋势,遗传传递力分别为110.48%、101.15%,110.2%;平均果形指数大于亲中值,与'玉露香'果形指数相接近;后代果皮颜色表现为4种,黄色、绿色、黄绿色与绿黄色,以黄色和绿色为主;杂交后代小果心的比率最大,占总数的56.4%;梨果点的遗传比较复杂,杂交后代出现比较明显的性状分离,果点明显的比率较大,占总数的44.7%。

内蒙古西南地区南果梨无公害栽培技术

南果梨(Pyrus ussuriensis‘Nanguo Pear’)是中国优良梨地方品种,营养丰富,风味好,适应性强。通过实地调查和多年经验积累,结合内蒙古西南地区南果梨的生长特点,提出了南果梨无公害栽培管理措施,以期为各地生产栽培南果梨提供借鉴。

梨果实品质评价因子的选择

测定了‘清香’、‘新雅’等26个梨(Pyrus pyrifolia)选育品种(系)的单果重、果形指数、果肉硬度、可滴定酸、可溶性固形物、可溶性糖、维生素C、糖酸比等8个主要品质因子,采用多元统计主成分分析法将上述8个相关的随机变量压缩成5个综合变量,再通过系统聚类分析,确定单果重、可溶性固形物、糖酸比、可滴定酸、果肉硬度为5个具有代表性的品质评价因子。