Effects of Low Temperature and Poor Lighting on Anthocyanin Content and Fruit Quality of Eggplant
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摘要:目的 探究低温、弱光、低温弱光处理对茄子幼苗期、花期、果期花青素含量的影响,以及对茄子品质的影响,为茄子的优质培育以及高产栽培奠定理论基础。方法 以紫黑茄秀娘为试验材料,分别在幼苗期、花期、果期进行低温(18 ℃/13 ℃,250 μmol·m−2·s−1)、弱光(25 ℃/20 ℃,120 μmol·m−2·s−1)、低温弱光(18 ℃/13 ℃,120 μmol·m−2·s−1)、CK(25 ℃/20 ℃,250 μmol·m−2·s−1)等4个处理,测定幼苗期形态及生理特性,不同时期、不同部位的花青素,以及果期果实的品质。结果 低温弱光胁迫对幼苗生长存在显著影响,在幼苗期低温对幼苗生长及生理影响显著大于弱光及低温弱光,花青素含量均表现为根<叶片<叶脉<茎;在花期,花青素含量依次为花萼<花瓣;在果期,花青素含量依次为果肉<果柄<果皮。茄子不同时期受到胁迫后,不同部位的花青素含量均呈现弱光<CK<低温弱光<低温,各胁迫下果实色泽指数依次为弱光<CK<低温弱光<低温,可溶性糖含量、可溶性蛋白含量、类黄酮含量、总酚含量均呈现低温<低温弱光<弱光<CK。结论 低温促进花青素合成;弱光抑制花青素合成;在低温弱光双因素互作下,低温因素对花青素含量的影响起主导作用,花青素的合成大于降解,花青素含量增加。低温、弱光、低温弱光胁迫下茄子品质均下降,其中,低温胁迫对茄子的品质影响最大。Abstract:Objective Anthocyanin content at various growth stages and fruit quality of eggplants exposed to low temperature and/or deficient light were studied.Methods Purple black eggplant Xiu Niang was grown in a greenhouse under (A) daytime/night temperatures of 18 ℃/13 ℃ with normal lighting at 250 μmol·m−2·s−1, (B) normal temperatures of 25 ℃/20 ℃ with poor lighting at 120 μmol·m−2·s−1, (C) low temperatures of 18 ℃/13 ℃ with poor lighting at 120 μmol·m−2·s−1, or (CK) normal temperatures and lighting. Growth, physiology, anthocyanin contents in different plant parts, and quality of fruit of the eggplants at seedling, flowering, and fruiting stages were monitored.Results The stresses of low temperature and/or poor lighting affected the growth of eggplant seedlings. Low temperature alone (i.e., A) exerted significantly greater effects on the growth and physiology of the seedlings than B or C. The anthocyanin contents in the organs of a seedling ranked stems>leaf veins>leaves>roots. At the flowering stage, the content was higher in the petals than the calyx; and at the fruiting stage, it ranked peels>stalks>fruits>pulp. The anthocyanin content in the plant at all stages under various treatments were B<CK<C<A. The coloration of eggplant was intensified by the treatments in a trend of B<CK<C<A. And the treatments appeared to cause reductions in the order of A<C<B<CK on the soluble sugars, soluble proteins, flavonoids, and total phenols contents in the plants.Conclusion Exposure to low temperature (e.g., 18 ℃ in daytime and 13 ℃ at night) promoted, but poor lighting inhibited, anthocyanin synthesis in eggplant. When both conditions were imposed simultaneously on the plants, the effect of low temperature on anthocyanin overshadowed that of poor lighting. In contrast, the fruit quality suffered by either low temperature, poor lighting, or both, especially low temperature.
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Keywords:
- low temperature /
- poor lighting /
- low temperature and poor lighting /
- growth stages /
- plant parts /
- anthocyanin /
- quality
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0. 引言
【研究意义】兔巴氏杆菌病是由多杀性巴氏杆菌感染兔引起的一种传染病。该病一年四季均可发生,且所有日龄的兔均可发病。临床上,兔巴氏杆菌病以呼吸道症状多见,也可见中耳炎和结膜炎[1-2]。该病是兔的常发病和多发病,是危害兔产业发展的重要疾病。根据多杀性巴氏杆菌的荚膜抗原可将其分为5种血清型,即A、B、D、E和F型[3]。其中,A和D型菌株是兔群中的优势流行菌株[4-5]。【前人研究进展】F型多杀性巴氏杆菌最早在美国的火鸡中发现[6],该菌株主要感染禽类[7],其感染能引起禽霍乱[8]。然而,研究表明国内外兔群中也存在F型多杀性巴氏杆菌,且该菌株对兔具有强致病性[9-11]。由此可见,兔群中F型多杀性巴氏杆菌的出现,使兔巴氏杆菌病的病因更加复杂,也使该病的确诊更加困难。因此,实现对F型多杀性巴氏杆菌的快速检测,掌握其在兔群中的流行情况,对兔产业的发展具有重要意义。【本研究切入点】目前,用于F型多杀性巴氏杆菌的实验室检测方法有细菌分离鉴定和多重PCR[3, 10]。多杀性巴氏杆菌营养需求高、生长较缓慢,容易受其他生长较快的细菌污染。多重PCR用于多杀性巴氏杆菌的荚膜分型[3],反应体系中包含6对引物,对反应体系的组成和反应条件都要求很高,否则会出现非特异性扩增或假阴性结果。【拟解决的关键问题】为了建立一种快速且简便的F型多杀性巴氏杆菌检测方法,本研究根据多杀性巴氏杆菌kmt1基因和F型多杀性巴氏杆菌fcbD基因的保守序列分别设计了2对特异性引物,建立了检测F型多杀性巴氏杆菌的双重PCR检测方法,为兔源F型多杀性巴氏杆菌的快速检测提供技术支撑。
1. 材料与方法
1.1 材料
1.1.1 主要试剂
2×PCR Mix、pEASY-T1克隆载体、细菌基因组DNA提取试剂盒和胶回收试剂盒购自北京全式金生物技术有限公司。
1.1.2 菌株
兔源A、D和F型多杀性巴氏杆菌(Pasteurella multocida)、支气管败血波氏杆菌(Bordetella bronchiseptica)、肺炎克雷伯菌(Klebsiella pneumonia)、大肠杆菌(Escherichia coli)、金黄色葡萄球菌(Staphylococcus aureus)由本实验室分离保存。
1.2 方法
1.2.1 引物设计
根据GenBank中公布的多杀性巴氏杆菌kmt1基因(登录号:KX348143)和F型多杀性巴氏杆菌的fcbD基因(登录号:AF302467),利用Primer Premier 5.0软件设计了2对分别针对kmt1基因和fcbD基因保守序列的特异性引物,kmt1基因引物序列为:kmt1-F:5′-gttttatgccacttgaaatgggaa-3′/kmt1-R:5′-taagaaacgtaactcaacatggaaatatt-3′;fcbD基因引物序列为:fcbD-F:5′-ctaaagatcttgttcttgctccattg-3′/fcbD-R:5′-tctgcggtaatattatgagtatccac-3′,扩增的目的片段分别为260 bp和490 bp。引物由上海铂尚生物技术有限公司合成。
1.2.2 单重PCR方法的建立及扩增产物的鉴定
以提取的兔源F型多杀性巴氏杆菌基因组DNA为模板,分别利用kmt1基因和fcbD基因引物进行单重PCR扩增。单重PCR反应体系为:2×PCR Mix 25 μL,基因组DNA 1 μL,上下游引物(10 μmol·L−1)各2 μL,灭菌ddH2O 20 μL,共50 μL反应体系。单重PCR反应程序为95 ℃ 5 min;95 ℃ 30 s、59 ℃ 30 s、72 ℃ 30 s,35个循环;72 ℃ 10 min。PCR产物纯化后克隆至pEASY-T1克隆载体,送上海铂尚生物技术有限公司测序。
1.2.3 双重PCR方法的建立及反应条件的优化
将kmt1基因和fcbD基因引物调整至40 μmol·L−1,等体积混匀后作为双重PCR的引物。双重PCR反应体系为:2×PCR Mix 25 μL,兔源F型多杀性巴氏杆菌基因组DNA 1 μL,混合引物4 μL,灭菌ddH2O 20 μL,共50 μL反应体系。双重PCR反应程序为95 ℃ 10 min;95 ℃ 30 s、59 ℃ 90 s、72 ℃ 2 min,35个循环;72 ℃ 10 min。在此基础上,设置双重PCR方法的退火温度在54~60 ℃、混合引物终浓度在0.4、0.5、0.6、0.7、0.8、0.9、1.0 μmol·L−1进行优化,确定最佳的退火温度和引物浓度。
1.2.4 双重PCR方法的特异性试验
分别以提取的兔源A、D和F型多杀性巴氏杆菌、支气管败血波氏杆菌、肺炎克雷伯菌、大肠杆菌、金黄色葡萄球菌的基因组DNA为模板,应用建立的双重PCR方法进行检测,设置阴性对照(灭菌ddH2O),评估该双重PCR方法的特异性。
1.2.5 双重PCR方法的敏感性试验
将荚膜F型多杀性巴氏杆菌的基因组DNA 10倍倍比稀释,使双重PCR反应体系中DNA模板的含量为1×107~1×100拷贝·μL−1,设置阴性对照(灭菌ddH2O),评估该方法的敏感性。
1.2.6 双重PCR方法的重复性试验
取90份已知结果的病死兔肺脏样品,平均分为3组,每组30份(A型多杀性巴氏杆菌样品5份,D型多杀性巴氏杆菌5份,F型多杀性巴氏杆菌5份,支气管败血波氏杆菌3份,肺炎克雷伯菌3份,大肠杆菌3份,金黄色葡萄球菌3份,阴性样品3份)。利用细菌基因组DNA提取试剂盒分别提取样品的基因组DNA,平均分为3份,利用建立的双重PCR方法分3次检测(每次检测90份),每次检测时每份样品重复3次,统计批内和批间变异系数,评估该双重PCR方法的重复性。
1.2.7 双重PCR方法的初步应用
选取从龙岩、三明、南平、福州和宁德5个地区收集的87份已知结果的呼吸道病死兔肺脏样品,应用本实验建立的双重PCR方法和已报道的多重PCR方法[3]同时对87份临床样品进行检测。统计检测结果,比较两种PCR方法检测结果与已知结果的一致性以及两种PCR方法检测结果的一致性。
2. 结果与分析
2.1 单重PCR方法的建立及扩增产物的鉴定
以兔源F型多杀性巴氏杆菌的基因组DNA为模板,利用设计的kmt1基因和fcbD基因引物分别进行单重PCR扩增。结果显示,扩增产物分别为260 bp和490 bp(图1),与预期目的片段大小相符。将上述2条目的片段克隆至pEASY-T1克隆载体并测序,测序结果显示2条目的片段序列与相应参考基因的序列同源性均为100%。
2.2 双重PCR方法的建立及反应条件的优化
以兔源F型多杀性巴氏杆菌的基因组DNA为模板,应用kmt1基因和fcbD基因引物进行双重PCR扩增。结果显示,在同一反应体系中,2对引物均能特异地扩增出相应的目的片段(图2)。在此基础上,进一步对双重PCR的退火温度和引物浓度进行优化。结果显示,当退火温度为54~60 ℃时,该双重PCR的扩增效果均较好(图3);当混合引物浓度为0.8 μmol·L−1时(图4),双重PCR扩增效果最好。退火温度高,则特异性强。因此,确定该双重PCR的最佳反应条件为退火温度60 ℃,混合引物浓度0.8 μmol·L−1。
2.3 双重PCR方法的特异性
利用建立的双重PCR能同时特异地扩增出兔源F型多杀性巴氏杆菌的kmt1基因片段和fcbD基因片段,能扩增出兔源A型和D型多杀性巴氏杆菌的kmt1基因片段,对兔源支气管败血波氏杆菌、肺炎克雷伯菌、大肠杆菌、金黄色葡萄球菌和阴性对照(灭菌ddH2O)则为阴性(图5)。结果表明,该双重PCR方法具有较强的特异性。
图 5 双重PCR检测方法特异性试验注:M: DNA Marker;1:F型多杀性巴氏杆菌;2:A型多杀性巴氏杆菌;3:D型多杀性巴氏杆菌;4:支气管败血波氏杆菌;5:肺炎克雷伯菌;6:大肠杆菌;7:金黄色葡萄球菌;8:阴性对照Figure 5. Specificity of duplex PCR assayNote: M: DNA marker; 1: serogroup F strain of P. multocida; 2: serogroup A strain of P. multocida; 3: serogroup D strain of P. multocida; 4:B. bronchiseptica; 5: K. pneumonia; 6: E. coli; 7: S. aureus; 8: negative control.2.4 双重PCR方法的敏感性
将兔源F型多杀性巴氏杆菌的基因组DNA 10倍倍比稀释(1×107~1×100拷贝·μL−1)。结果显示,该双重PCR的最低检测限为1×103拷贝·μL−1基因组DNA(图6),表明该双重PCR具有良好的敏感性。
2.5 双重PCR方法的重复性
应用建立的双重PCR对90份已知结果的病死兔肺脏样品(3组,每组30份)分3批次进行批内和批间重复性试验。结果显示,重复性试验批内和批间结果均一致,表明该双重PCR具有良好的重复性。
2.6 双重PCR方法的初步应用
应用建立的双重PCR方法和已报道的多重PCR方法同时对87份已知结果(A型多杀性巴氏杆菌阳性样品30份,D型多杀性巴氏杆菌阳性样品9份,F型多杀性巴氏杆菌阳性样品8份,支气管败血波氏杆菌阳性样品11份,肺炎克雷伯菌阳性样品2份,大肠杆菌阳性样品1份,金黄色葡萄球菌阳性样品3份,阴性样品23份)的呼吸道病死兔肺脏样品进行检测。结果显示,双重PCR检测出多杀性巴氏杆菌阳性样品49份(其中F型多杀性巴氏杆菌阳性样品8份),阴性样品38份。多重PCR检测出多杀性巴氏杆菌阳性样品43份(其中A型多杀性巴氏杆菌阳性样品30份,D型多杀性阳性样品7份,F型多杀性巴氏杆菌阳性样品6份),阴性样品39份,非特异扩增样品5份。双重PCR方法检测结果和已报道的多重PCR方法检测结果与已知结果的符合率分别为97.70%和94.25%。双重PCR方法检测结果与已报道的多重PCR方法检测结果的符合率为93.10%。上述结果表明,本试验建立的双重PCR方法准确性高,具有较好的临床应用价值。
3. 讨论与结论
多杀性巴氏杆菌感染是引起兔呼吸道疾病的重要病原之一,常常引起致死性感染。临床上,致死性病例以50~70日龄的商品兔、怀孕后期母兔和哺乳母兔多见,给养兔业造成严重的经济损失[12]。兔巴氏杆菌病主要由A和D型多杀性巴氏杆菌感染引起[4-5]。F型多杀性巴氏杆菌首次分离自火鸡[6],主要在禽类中流行病且致病性强[7-8]。然而,在国内外兔群中也发现有该菌的存在,且其感染能引起兔的严重致死性呼吸道疾病[9-11]。由此可见,F型多杀性巴氏杆菌在兔群中的出现使兔巴氏杆菌病病因更加复杂,导致该病的确诊更加困难。
本试验根据多杀性巴氏杆菌的kmt1基因和F型多杀性巴氏杆菌fcbD基因的保守序列分别设计了2对特异性引物,建立了检测F型多杀性巴氏杆菌的双重PCR方法。kmt1基因是多杀性巴氏杆菌的种特异性基因,以该基因为目的基因能建立检测多杀性巴氏杆菌的特异性PCR检测方法[13-14]。fcbD基因编码F型多杀性巴氏杆菌荚膜中的软骨素,是F型菌株中的特异性基因,以该基因为目的基因能建立鉴定F型多杀性巴氏杆菌荚膜血清型的多重PCR方法[3]。由此可见,以kmt1基因和fcbD基因为目的基因能建立特异的检测兔源F型多杀性巴氏杆菌的双重PCR检测方法。本试验建立的双重PCR方法快速简便,不仅克服了细菌分离鉴定的费时,还克服了多杀性巴氏杆菌荚膜分型多重PCR方法的费力。此外,该双重PCR方法特异性强、重复性好、准确性高,具有很好的临床应用价值,为掌握兔群中F型多杀性巴氏杆菌的流行情况提供了有力的技术手段。
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表 1 低温、弱光、低温弱光胁迫对茄子幼苗形态及生理特性的影响
Table 1 Effects of Low temperature and poor lighting on growth and physiology of eggplant seedlings
处理
Treatment叶面积
Leaf
area/
mm2根表
面积
Root
surface area/
mm2叶绿素
含量
Chlorophyll
content/
(mg·g−1)Fm' ΦPSII NPQ qp 可溶性蛋白
含量
Soluble
protein
content/
(mg·g−1)可溶性糖
含量
Soluble
sugar
content/
(mg·g−1)丙二醛
含量
MDA/
(mg·g−1)相对电导率
Relative
conductivity/%CK 139.9±15.48b 78.05±2.34a 1.56±0.06a 0.74±0.03a 0.45±0.01a 0.20±0.01c 0.73±0.03b 4.47±0.38c 1.71±0.09c 0.21±0.01c 0.68±0.03 d S1 105.86±10.70c 59.39±2.16c 1.30±0.01c 0.63±0.02b 0.40±0.01b 0.29±0.01a 0.66±0.01c 6.99±0.91a 3.32±0.32a 0.30±0.01a 0.94±0.01a S3 233.69±26.54a 59.00±14.03c 1.21±0.05d 0.59±0.03c 0.33±0.03c 0.29±0.01a 0.57±0.05d 5.72±0.30b 2.11±0.08b 0.28±0.00ab 0.89±0.00b S5 143.78±5.42b 72.02±2.09b 1.41±0.03b 0.62±0.02b 0.46±0.02a 0.26±0.01b 0.79±0.03a 4.82±0.43c 2.15±0.24b 0.26±0.00b 0.80±0.01c 表 2 低温、弱光、低温弱光胁迫下茄子的色泽指数
Table 2 Color indexes of eggplant under various treatments
胁迫
Treatment亮度
Lightness L红色度
Redness a黄色度
Yellowness b颜色指数
CIRGCK 26.74±1.81b 3.72±0.89b 0.15±0.04b 5.93±0.48c S1 23.07±1.03c 1.99±0.32c −1.25±0.30 d 7.11±0.19a S3 34.65±1.21a 6.88±0.57a 2.69±0.46a 4.27±0.13b S5 23.64±0.57c 2.81±0.77bc −0.38±0.04c 6.81±0.19a 表 3 低温、弱光、低温弱光胁迫对茄子果实品质的影响
Table 3 Effects of various treatments on eggplant fruit quality
胁迫
Treatment可溶性糖含量
Soluble sugar content/%可溶性蛋白含量
Soluble protein content/(mg·g−1)类黄酮含量
Flavonoid content/(mg.g−1)总酚含量
Total phenol content/(mg.g-1)CK 8.37±150.48a 14.86±0.14a 0.58±0.02a 2.22±0.04a S1 5.96±150.17 d 9.59±0.51 d 0.31±0.03 d 0.98±0.20 d S3 7.66±150.10b 11.73±0.54b 0.45±0.05b 1.85±0.01b S5 6.86±150.43c 10.38±0.14c 0.40±0.02c 1.51±0.07c -
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