摘要
西瓜(Citrulls.lanatus)在整个果实发育期都存在裂果现象,导致产量和质量降低,也不利于长距离运输,伤口容易产生病害,生产成本增加,开展西瓜裂果研究亟不可待。目前,关于西瓜裂果的机制研究报道很少,采用什么标准评价西瓜裂果程度,西瓜裂果的遗传机制还不明确,一些优良抗裂西瓜资源未得到很好利用。本研究进行一组试验,旨在从形态学观察、理化机理、遗传机制、分子标记筛选等多方面、多层次探求西瓜裂果机制,提出西瓜抗裂性的评价标准和单株裂果鉴定技术,将传统研究方法和分子技术相结合,探索西瓜裂果相关关键理论和应用问题,为进一步开展西瓜种质改良和抗裂育种奠定基础。主要研究成果如下：
1研究西瓜果实性状与田间抗裂果性的相关性、西瓜抗裂性评价标准以及西瓜单株裂果鉴定技术体系
在普通大棚和露地两种栽培条件下鉴定西瓜裂果性,制定西瓜的裂果性状分级标准和抗性评价标准；不同基因型西瓜裂果性差异显著(F保护地=10.7218**, F露地=5.783**,F0.01=1.90,df=13),大棚和露地两种栽培条件下抗裂程度显著相关(r=0.937**,r 0.01=0.6614,df=13)。
对西瓜的14个农艺性状进行相关与通径分析结果表明裂果率与外果皮绿皮层呈显著相关,可将其作为筛选抗裂品种的重要参考性状。
以田间裂果率为参照指标,采用相关显著性分析和聚类分析等方法,比较了形态指标鉴定(果皮厚度、外果皮绿皮层厚度)、人工诱导鉴定(耐压法、浸液减压法、切裂应度法)等方法的稳定性、可靠性,结果表明,果实切裂应度法,测定简单方便、受客观环境的影响相对较小,是西瓜单株裂果鉴定较佳方法。
2水分、钾肥处理对西瓜裂果的影响
通过采取控制水分和漫灌两个处理,测定裂果和未裂果西瓜果实各部分含水量,结果表明控水处理的西瓜裂果率显著降低；西瓜田间裂果率与裂果果皮含水量极显著负相关(r=-0.975**),与果瓤果皮含水量差异比值显著正相关(r=0.9074*),与裂果果瓤和未裂果果瓤含水量相关性不显著。
以苏密1号、京欣1号西瓜品种以及筛选的不同裂果差异性的自交系为材料,研究了不同浓度的KCl叶面喷施对西瓜果实裂果性及几种生理保护性酶的影响,结果表明,超氧化物歧化酶(SOD)、过氧化物酶(POD)和丙二醛(MDA)与田间裂果率的简单相关系数分别为-0.52843、-0.46383、0.15047,均未达到显著相关,叶面喷施0.1%、0.2%、0.4%、0.6%KCl,随着浓度的增加,裂果率降低,SOD、POD的含量增加,MDA的量降低。
3西瓜组合X-10l×L-103果实切裂应度、果皮绿皮层厚度的遗传模型和遗传参数
应用植物数量性状主基因+多基因混合遗传模型多世代联合分析方法,对组合L-103×X-101的果实切裂应度、果皮绿皮层厚度进行遗传分析。结果表明：L-103×X-101组合果实切裂应度性状的遗传受2对加-显-上位性主基因+加-显多基因控制；B1、B2、F2群体的主基因遗传率分别为40.34%、25.08%、33.10%,而多基因遗传率遗传率为32.65%、0、50.20%；在组合后代群体中遗传变异平均占表型变异的60.45%,各群体的环境平均变异率达39.55%,环境对西瓜果实裂果性状有较大影响。果皮绿皮层厚度的遗传受2对加性-显性-上位性主基因+加性-显性多基因控制, B1、F2群体的主基因遗传率分别为94.63%、93.88%,而B2群体的主基因遗传率为0,多基因仅在B1中有遗传,且遗传率很低,仅为0.544%,果皮绿皮层厚度受环境变异的影响非常小。在育种实践中,我们可以利用其高遗传效率,进行早期选择。
4建立西瓜SRAP技术体系,筛选出与西瓜抗裂性状连锁的SRAP标记
利用正交试验设计,对SRAP-PCR反应体系中的Mg2+浓度、dNTps浓度、引物浓度、Taq酶浓度和模板DNA浓度进行5因素4水平的筛选分析,确定西瓜SRAP-PCR最佳反应体系为：10xPCRbuffer2uL、Mg2+3.0mmol/L, dNTPs 0.2mmol/L,引物0.5umol/L,模板DNA40ng, Taq聚合酶0.5U,总体积为10uL。
利用优化的SRAP体系和300对SRAP引物在P1、P2、极端抗裂池、极端感裂池的DNA进行扩增筛选引物,引物me8-em1在父本和抗裂池的300bp-400bp之间扩增出稳定、清晰地条带,而母本和感裂池没有条带,可能是与裂果相关的SRAP标记。在163个F2单株中验证,结果显示,在138个抗裂F2单株中126个单株产生了与抗裂亲本相同的带型,25个表现感裂的单株仅有5株产生了与抗裂亲本相同的带型,利用Mapmaker version 3.0计算出该标记与抗裂性状的连锁距离为9.9cM。
The fruits of watermelon are often found cracking duiring the whole Fruit development. Fruit-cracking results in lower yield and quality, easier infection to the deseases and more difficulties for sales. So it is a great significance to study mechanism of fruit-cracking in watermelon. However, fruit-cracking indicators and evaluation criteria have not yet established, and some good crack resistance merterials have not been well used. Therefore, a group of tests, such as morphology, physical and chemical mechanisms, genetic mechanism, and molecular marker and so on, was carried out for understanding watermelon fruit cracking. In this study, we proposed evaluation criteria and reliable identification technology on fruit cracking-risistance. On these bases, we combine traditional methods with molecular technology for in-depth exploration fruit cracking theory and application. It is very important for further breeding on crack-resistant and germplasm improvement in watermelon. The major results are as follows:
1. The correlation between fruit agronomic traits and fruit-cracking of watermelon, grades and symptoms of fruit-cracking in the field, and an accurate, stable, scientific identification method of fruit cracking were studied.
Resistance to fruit cracking of different gene-types watermelon was evaluated under opening and protected conditions. Grading standards and evaluation criteria of fruit-cracking in watermelon are drawn up. According to the comprehensive evaluation, the variance of the cracking indices were significant in opening field (F=5.783**, F0.01=1.90, df=13) and protected field (F=10.7218**, F0.01= 1.90, df=13). The correlation coefficients of cracking indices in both conditions were significant (r=0.937**, r0.01=0.6614, df=13).
The correlation and path analysis on 14 agronomic traits of watermelon were studied. The thickness of pericarp and green epicarps were significantly related to the rate of fruit cracking and can be taken as a main reference trait for selecting anti-cracking varieties.
By taking fruit-cracking rate of inbred line in the field as reference, we compare the stability, sensitivity and practicality of different identification methods, such as morphological index (pericarp thickness, green pericarp) and artificially inducing methods (pressure method, extract decompression, cut cracking), by correlation analysis and cluster analysis. The results show that the fruit cut cracking method, which is easy and convenient to determinate and is practicable to reflect the true fruit resistance of watermelon varieties, can be used as the identification method of fruit cracking in watermelon.
2. The relationship between water, KC1 and rate of fruit cracking in watermelon
The impact of water on watermelon fruit cracking is discussed. In the condition of the Water-control and water irrigation, we determinate water content of the various parts of cracking fruit and non-crack fruit. The results shows, watermelon fruit cracking rate, in water-controlled treatment, was significantly reduced; The relationship between rate of fruit cracking and water content in the rind of fruit cracking was significantly negatively correlated (r=-0.975), and the ratio of water content of fruit pulp and peel between cracking and non-cracking fruit was significantly correlated (r= 0.9074).
The relation among KCl, physiological Emezy and fruit cracking resistance of watermelon were studied. The results show, correlation coefficient between superoxide dismutase (SOD), peroxidase (POD), malondialdehyde (MDA) and fruit cracking rate in the field was -0.52843,-0.46383, and 0.15047, which have no significant variance. When we spray KCl on the leaves during the development of watermelon, the cracking rate is decreasing, SOD, POD are increasing and MDA decreasing with the KCl concentration increased.
3. Genetic Analysis of Fruit cut cracking and Green epicarpin Watermelon
In this paper, inheritance of fruit cut cracking and green epicarpwere analyzed by applying the joint segregation analysis of a mixed genetic model of major gene plus polygene in six generations(P 1, P2, F1, B1, B2, and F2)derived from two crosses of L-103 (fruit-cracking susceptible)×X-101 (fruit-cracking resistance). The cut fruit cracking was dominated by two major genes with additive-dominance-epistatic effects plus polygenes with additive-dominance effects(E-1 model)in Cross X-101×L-103, The hereditability of major genes(hmg2)was 40.34%、25.08%、33.10% in B1、B2、F2 population, and that of polygenes(hpg2) detected in B1、B2、F2population was 32.65%、0、50.20%. It also indicated that the environmental variance and the genetic variance accounted for 39.55% and 60.45%, on everage, of the phenotypic variance in each population respectively, indicating that the environmental factors have a great effect on fruit cracking resistance. The green epicarpwas dominated by two major genes with additive-dominance-epistatic effects plus polygenes with additive-dominance effects (E-1 model) in the Cross. Hereditability of major genes were 94.63%、93.88% in F2 and Blpopulation, while was zero in B2 population. Hereditability of polygenes only detected in B1 population was 0.544%. It indicated that the green epicarpin is dominated by major genes. In addition, the environmental variance was little in F2 population, so hereditability of major genes can be favorable to an efficient selection in early generation.
4. The SRAP reaction system of Watermelon and screen of SRAP molecular markers linked with fruit cracking resistance
In this experiment, the reaction system of watemelon SRAP was optimized, in terms of Mg2+, dNTPs, primers, Taq E and template DNA 5 factors and their 4 levels with the primer Em3-Me3, by an orthogonal design.The watermelon appropriate SRAP-PCR reaction system in a 10μL reaction system was:10xPCRbuffer2uL, Mg2+3.0 mmol/L, dNTPs-0.2mmol/L, primer 0.5umol/L, template DNA40ng, Taq polymerase 0.5U.
In this paper, bulked segregant analysis was used to screen SRAP molecular markers linked to fruit anti-cracking of watermelon using two P1, P2 and extreme types of DNA pools. By the watermelon appropriate SRAP-PCR reaction system, 1(me8-eml) primer was found with polymorphic bands from 300 pairs of SRAP primer combinations. These were further amplified in 163 F2 single DNA. The results shows 125 single plant DNA have the same maker band as the anti-cracking parent in 138 resistant single plant, Only 5 single plant DNA has the maker in 25 susceptible single plant. This marker was testified with individual DNA of the F2 population and the band could only be amplified in the high cracking resistant plants. Linkage analysis using the software of MAPMAKER/Exp3.0 indicated its genetic distance to the fruit cracking resistance is 9.9cM.
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