钙对SA诱导番茄抗灰霉病的作用机制研究
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摘要
番茄(Solanum lycopersicum Mill)是多发病的重要园艺作物。目前，生产者防治其病害的重要方法之一就是依赖农药，这对环境和农产品造成严重污染是不言而喻的。诱导抗性是植物自身所产生的一种更为主动的抗性反应方式，研究植物诱导抗性具有重要的理论意义和实际应用价值。
钙(Ca~(2+))和水杨酸(SA)均被认为是重要的抗逆性物质（Sugimoto等，2005; Volpinand Elad,1991; Yoon CS等，2010; White,1979; Durrant2004; Mauch2001; Hayat andAhmad,2007），特别是SA被认为是重要的植物诱导抗病物质（White,1979; Durrant2004;Mauch2001; Hayat and Ahmad,2007）。然而，尽管多项研究已表明钙(Ca~(2+))在SA诱导的植物ROS爆发、耐热性、抗旱性、气孔开闭、采后生理等方面均有调控作用（Pei等，2000；刘悦萍等，2005；王利军等，2003；刘新等，2003；敖日嘎，2003），但钙(Ca~(2+))对SA诱导植物抗病的作用尚缺乏详细报道，更缺乏对其机制的探讨。
本研究以番茄灰霉病敏感型品种“L402”为试材，从生理及分子水平上分析了Ca~(2+)对SA诱导番茄抗灰霉病的作用机制。以期为利用Ca~(2+)和SA增强番茄植株抗病性、提高番茄产量和品质奠定理论和应用基础。主要研究结果如下：
1.明确了营养液中增施Ca~(2+)和叶面增施Ca~(2+)均显著降低了番茄灰霉病的病情指数，说明Ca~(2+)具有提高番茄植株抗灰霉病的能力，其中营养液中增施Ca~(2+)较对照病情指数降低63.61%，叶面增施Ca~(2+)较对照降低6.04%，说明根际增施Ca~(2+)效果更显著；但叶面增施Mg~(2+)则显著提高番茄灰霉病的病情指数，病情指数达78.25，比对照提高4.54%，表明镁(Mg~(2+))具有降低番茄抗灰霉病的作用。
2.针对根际增施Ca~(2+)抗病效果显著做了进一步分析，明确了营养液Ca~(2+)浓度处理通过PAL合成途径提高番茄植株叶片内源SA含量。PAL合成途径CM，PAL基因表达水平和BA2H酶活性在营养液Ca~(2+)浓度处理条件下显著提高，但对增强PAD4和ICS1基因表达的作用不显著，因此认为Ca~(2+)通过调控PAL途径提高番茄植株叶片内源SA含量。
3.明确了叶面增施SA显著降低了番茄灰霉病的病情指数；Mg~(2+)与SA配施显著降低了SA诱导番茄抗灰霉病的作用。而Ca~(2+)与SA不同顺序配施对番茄幼苗抗灰霉病的影响效果不同，先施Ca~(2+)再施SA显著降低了番茄灰霉病的病情指数，即Ca+SA处理的病情指数比SA降低17.93%，比SA+Ca降低13.34%，比单纯施Ca~(2+)降低45.22%；而SA+Ca处理的病情指数与SA处理无显著差异。说明先施Ca~(2+)再施SA，Ca~(2+)具有显著增强SA诱导番茄抗灰霉病的作用。
4.明确了Ca~(2+)、Ca~(2+)抑制剂EGTA与SA单独或配合施用在诱导及接种条件下对番茄植株叶片ROS积累、PAL、几丁质酶和β-1,3-葡聚糖酶活性及基因表达的影响。其中SA处理诱导了番茄植株叶片的ROS爆发，Ca~(2+)促进了SA诱导的番茄植株叶片ROS爆发；EGTA抑制了ROS积累，而且抑制了SA诱导的ROS积累。Ca~(2+)及SA对植株抗病相关酶PAL、几丁质酶和β-1,3-葡聚糖酶活性及其基因表达的影响同ROS爆发具有相似的结果。说明SA可能通过调控植株抗病相关酶的基因表达及ROS爆发进而提高植株抗灰霉病能力的，而Ca~(2+)通过促进SA诱导植株抗病相关酶基因表达及ROS爆发而进一步提高植株抗灰霉病能力。
5.明确了Ca~(2+)对SA诱导番茄抗灰霉病蛋白表达的调控作用。采用TCA/丙酮法提取各处理接种灰霉菌48h后番茄幼苗叶片的蛋白，并采用双向电泳技术及PD-Quest软件分析，发现每张图谱可检测到超过500个蛋白点，软件分析得出65个差异蛋白点，在两个处理中均表达上调的蛋白有19个；两个处理都表达下降的蛋白点有6个；两个处理均是新出现的蛋白点有9个；此外，在SA处理中，有2个蛋白点仅在该处理中呈上调表达，有3个蛋白点仅在该处理中呈下调表达，有2个蛋白点仅在该处理中是新出现的蛋白点；有2个蛋白点在该处理中是消失的蛋白点。在Ca+SA处理中，有8个蛋白点仅在该处理中呈上调表达，有1个蛋白点在该处理中呈下调表达，10个蛋白点仅在Ca+SA处理中是新出现的蛋白点，有3个蛋白点仅在该处理中表达消失。
6.采用基质辅助激光解析电离飞行时间质谱技术(MODI-TOF-TOF)将稳定表达、丰度值变化1.5倍以上的蛋白点，并根据以往经验排除功能性蛋白，挑选出其中的50个进行鉴定，根据NCBI数据库搜索比对，鉴定成功了42个蛋白质点。它们分别参与了植物自身的防卫反应、能量代谢、蛋白质合成和转录调控等多个生理生化过程。明确了病程相关蛋白5和STH2、酸性内源几丁质酶等抗病相关物质以及SOD、APX等ROS相关酶在Ca~(2+)促进SA诱导番茄抗病性中起到了关键性的作用。
7.营养液Ca~(2+)浓度处理配合施用SA对NahG及其野生型（Moneymaker）的病情指数调查表明，在SA存在条件下Ca~(2+)可显著提高番茄抗病性。分析了SA依赖的信号途径下游关键基因(NPR1、PR1、TGA1a、TGA2.2)及蛋白激酶基因（MPK2、MPK4）的表达情况。NPR1、PR1、TGA1a、TGA2.2在补Ca~(2+)处理中表达均明显上调，而MPK4则表达趋势与此相反，仅在0mM Ca~(2+)处理中表达才上调，而0mM Ca~(2+)处理番茄抗病性显著降低。证明了SA信号转导相关基因及MPK2基因表达上调是营养液Ca~(2+)浓度处理配合施用SA提高番茄抗病性的主要原因，且与MPK4无关。
8.明确了营养液Ca~(2+)浓度处理再增施SA对Ca~(2+)受体相关基因表达的影响。与3mMCa~(2+)处理相比8mM Ca~(2+)处理明显提高了CDPK的表达水平，0mM Ca~(2+)处理中则明显下降；CaM的表达显示其对外源SA处理较为敏感；CBL的上调表达可能受Ca~(2+)和SA的共同作用。因此推测营养液Ca~(2+)配合SA处理提高番茄抗灰霉病可能部分通过Ca~(2+)受体调节SA信号途径相关基因表达。
9.明确了营养液Ca~(2+)处理均可显著降低野生型‘Moneymaker’和SA缺失突变体‘NahG’品种的病情指数，但野生型‘Moneymaker’比SA缺失突变体‘NahG’的病情指数更低；Ca+SA处理进一步显著降低番茄野生型‘Moneymaker’和SA缺失突变体‘NahG’品种的病情指数，同样野生型‘Moneymaker’比SA缺失突变体‘NahG’的病情指数更低；说明Ca~(2+)既有与SA共同降低番茄病情指数的作用，Ca~(2+)也具有不依赖SA降低番茄病情指数的作用，详细机制有待进一步研究。
Tomao (Solanum lycopersicum Mill) is an important kind in horticulture production inthe world and China. However, tomato is also a frequently-occurring crop. Currently, theproducer mainly depended on the chemicals to prevent disease，which lead to environmentaland products pollution. The induced resistance is an action mode that comes from plantsthemselves with the characteristic of activity. And which was worth to be researching andapplication. We should carry on depth research on the mechanism of induced resistance if wewant to exploit the plant induce resistance in order to enhance plant defence.
Ca~(2+)and Salicylic acid are considered that the important resistance substance (Sugimotoet al.,2005; Volpin and Elad,1991; Yoon CS et al.,2010; White,1979; Malamy et al.,1990;Gaffney et al.,1993; Durrant2004; Mauch2001). A growing number of detail reports indicatethat Ca~(2+)have strength role on SA induced effects, such as ROS, sub-high temperatureresistance, drought resistance, on-off of stomata and pos-harvest physiology of peach (Pei etal.,2000；Liu et al.,2005; Wang et al.,2003; Liu et al.,2003; Ao,2003). There is not detailreport on the mechanism of Ca~(2+)effect on SA induced resistance on Botrytis cinerea infectedtomato, and research of the mechanism is lack.
The cultivated tomato ‘L402’ which was sensitive to Botrytis cinerea, was employed inthis experiment, the treatments of with or without Ca~(2+)was designed, base on the level ofphysiology and molecule analysised the Ca~(2+)effect on SA induced resistance on Botrytiscinerea inoculation. It is helpful to study the mechanism to achieve the finally objective toenhanced the SA resistance efficiency and the yield and qulity both in scientific and practicalfields. The main research results are as follows:
1. It was clear that Ca~(2+)have the ablity to enhanced plants resistance through anaylisis thedisease index of different Ca~(2+)concentration of nutrient solution and foliar application Ca~(2+).Compare with control, the disease index declined63.61%in the treatment of supplementingCa~(2+)to nutrient solution; and the disease index reduced6.04%in treatment of foliarapplication Ca~(2+)compare to control; which illustrated that it was efficiency to apply Ca~(2+)tothizosphere. It was found that Magnesium (Mg~(2+)) did negative effect on tomato resistance onBotrytis cinerea, in alone Mg~(2+)treatment the disease index was78.25，which was increase4.54%compares with control.
2. Aimed to analysis the detail reasons of effectively enhancing the tomato resistanceunder thizosphere Ca~(2+)supplementation. It was found that the effect of Ca~(2+)treatment on SAaccumulation and SA synthesis in tomato leaves. It was showed that the expression level ofCM、PAL and activity of BA2H increased significantly Ca~(2+)treated, however, the expressionlevel of PAD4and ICS1were not significantly. So SA accumulation in Ca~(2+)concentrationtreated, which was the results of PAL pathway related genes expression levels’ increase.
3. It was clear that the disease index reduced significantly by foliar appling SA; andMg~(2+)did negative effect on SA induced resistance on Botrytis cinerea. The efficiency was difference that Ca~(2+)and SA in different order were applied to test the resistance on Botrytiscinerea. The Ca+SA treatment have the best effect on defense, the disease index decrease17.93%compare with alone SA treatment, and13.34%compare with SA+Ca, there was notdifference between SA+Ca and SA treated. The Mg~(2+)declined SA-induced resistance on graymold; the results of application of SA after Ca~(2+)indicated that Ca~(2+)enhanced SA-inducedresistance on gray mold in tomato.
4. It was found that Ca~(2+), EGTA, SA alone and applied complex led to ROS, enzymesactivity and genes expression increase under induced treatments or inoculation with graymold after induced treatments. Among the different treatments, the results showed SA caninduce ROS and Ca~(2+)enhanced the SA-induced ROS. In contrast，pretreatment with Ca~(2+)chelator EGTA, weakened the ROS and weakened the SA-induced ROS. The similar resultswere found in PAL, chintase, β-1,3-glucancse activity and genes expressions (PR1、 PR2aand PR3b). It indicated that SA enhanced tomato resistance by regulation of ROS andpathogensis related protein activity and expression, and Ca~(2+)promoted SA-induced resistanceto tomato gray mold by further increase these index.
5. It was clear that the function of proteins expression of Ca~(2+)regulating SA-inducedresistance to gray mold. The tomato leaves protein was extracted with TCA/acetone under theCK、SA、Ca+SA treatments inoculation with gray mold after48h. There were more than500protein spots in gel image through2-DE technical and the software of PD-Quest. We totallyfound out65differentially expressed protein spots. In detail, there were19spots wup-regulated,6spots down-regulated,9new spots in SA and Ca+SA treatemts; in addition,2spots up-regulated,3spots down-regulated,2new spots and2spots disappeared only in SAtreatment; and8spots up-regulated,1spots down-regulated,10new spots and3spotsdisappeared only in Ca+SA treatemts.
6.50differentially expressed protein spots with stable expression and higher than1.5-fold spot density were analysed by MODI-TOF-TOF. The42protein spots were obtainedby NCBI database searehing and intereomparison preliminary identifieation. These proteinswere associated with a variety of functions, including photosynthesis, energy metabolism,metabolism, ROS, protein synthesis and degradation, cell growth/division, signal transduction,and transcriptional. The key protein and enzyme of Ca~(2+)enhanced SA-induced resistance ontomato Botrytis cinerea were pathogenesis-related protein5, PR STH-2, acidic endochitinase,SOD and APX.
8. It was found that the disease index of treatments of Ca~(2+)concentration of nutrientsolution with SA application reduced significantly in ‘NahG’ and WT, which proved that Ca~(2+)enhanced tomato resistance at the condition of SA presence. Effects of Ca~(2+)on SA signalingtransduction in tomato was observed in this paper. The SA-dependent signaling related keygene (NPR1、PR1、TGA1a、TGA2.2) and two mitogen-activated protein kinase (MPK2、MPK4)expression level were analysised. An obvious increase of NPR1、PR1、TGA1a、TGA2.2、MPK2transcript was observed in8mM Ca~(2+)treated tomato seedlings. In contrary, the MPK4 expression level was up-regulation only in0mM Ca~(2+)treated, and the resistance declinedsignificantly in the treatment of0mM Ca~(2+). Looking From the rule of changes, Ca~(2+)concentration of nutrient solution with SA application enhanced tomato resistance, which wasresults of up-regulation expression of SA signaling pathway related genes and MPK2, not theMPK4.
9. The gene expression of Ca~(2+)related receptor had a great change after treatment ofCa~(2+)concentration of nutrient solution with SA application. The transcription of CDPKenhanced obviously in8mM Ca~(2+)comparation with3mM Ca~(2+), which expression level waslower in0mM Ca~(2+)than3mM Ca~(2+)treatment. From the analysis of relationship betweenCa~(2+)concentration and CaM expression level, it was indicated that the CaM transcription waspositively correlated to SA simulation. The most probably of CBL transcription situation wasCa~(2+)and SA co-regulation. So we speculated that Ca~(2+)related receptor partly participated inthe process of Ca~(2+)concentration with SA application treatment to enhance tomato resistance.
7. It was clear that Ca~(2+)have the ablity to enhanced plants resistance through treated‘Moneymaker’ and’NahG’ with different Ca~(2+)concentration of nutrient solution andanaylisis the disease index. The disease was lower in ‘Moneymaker’comparation to ‘NahG’.Further more, the disease index was lower in ‘Moneymaker’ and’NahG’ treated with2mMSA after appling with different Ca~(2+)concentration of nutrient solution comparation totreatments of different Ca~(2+)concentration of nutrient solution, it showed that Ca~(2+)not onlyhad the ablity to reduce tomato disease index combined with SA, but also had the function toinclined tomato disease index by the mode of SA-independent. The detail mechanism need tofurther research.
钙(Ca~(2+))和水杨酸(SA)均被认为是重要的抗逆性物质（Sugimoto等，2005; Volpinand Elad,1991; Yoon CS等，2010; White,1979; Durrant2004; Mauch2001; Hayat andAhmad,2007），特别是SA被认为是重要的植物诱导抗病物质（White,1979; Durrant2004;Mauch2001; Hayat and Ahmad,2007）。然而，尽管多项研究已表明钙(Ca~(2+))在SA诱导的植物ROS爆发、耐热性、抗旱性、气孔开闭、采后生理等方面均有调控作用（Pei等，2000；刘悦萍等，2005；王利军等，2003；刘新等，2003；敖日嘎，2003），但钙(Ca~(2+))对SA诱导植物抗病的作用尚缺乏详细报道，更缺乏对其机制的探讨。
本研究以番茄灰霉病敏感型品种“L402”为试材，从生理及分子水平上分析了Ca~(2+)对SA诱导番茄抗灰霉病的作用机制。以期为利用Ca~(2+)和SA增强番茄植株抗病性、提高番茄产量和品质奠定理论和应用基础。主要研究结果如下：
1.明确了营养液中增施Ca~(2+)和叶面增施Ca~(2+)均显著降低了番茄灰霉病的病情指数，说明Ca~(2+)具有提高番茄植株抗灰霉病的能力，其中营养液中增施Ca~(2+)较对照病情指数降低63.61%，叶面增施Ca~(2+)较对照降低6.04%，说明根际增施Ca~(2+)效果更显著；但叶面增施Mg~(2+)则显著提高番茄灰霉病的病情指数，病情指数达78.25，比对照提高4.54%，表明镁(Mg~(2+))具有降低番茄抗灰霉病的作用。
2.针对根际增施Ca~(2+)抗病效果显著做了进一步分析，明确了营养液Ca~(2+)浓度处理通过PAL合成途径提高番茄植株叶片内源SA含量。PAL合成途径CM，PAL基因表达水平和BA2H酶活性在营养液Ca~(2+)浓度处理条件下显著提高，但对增强PAD4和ICS1基因表达的作用不显著，因此认为Ca~(2+)通过调控PAL途径提高番茄植株叶片内源SA含量。
3.明确了叶面增施SA显著降低了番茄灰霉病的病情指数；Mg~(2+)与SA配施显著降低了SA诱导番茄抗灰霉病的作用。而Ca~(2+)与SA不同顺序配施对番茄幼苗抗灰霉病的影响效果不同，先施Ca~(2+)再施SA显著降低了番茄灰霉病的病情指数，即Ca+SA处理的病情指数比SA降低17.93%，比SA+Ca降低13.34%，比单纯施Ca~(2+)降低45.22%；而SA+Ca处理的病情指数与SA处理无显著差异。说明先施Ca~(2+)再施SA，Ca~(2+)具有显著增强SA诱导番茄抗灰霉病的作用。
4.明确了Ca~(2+)、Ca~(2+)抑制剂EGTA与SA单独或配合施用在诱导及接种条件下对番茄植株叶片ROS积累、PAL、几丁质酶和β-1,3-葡聚糖酶活性及基因表达的影响。其中SA处理诱导了番茄植株叶片的ROS爆发，Ca~(2+)促进了SA诱导的番茄植株叶片ROS爆发；EGTA抑制了ROS积累，而且抑制了SA诱导的ROS积累。Ca~(2+)及SA对植株抗病相关酶PAL、几丁质酶和β-1,3-葡聚糖酶活性及其基因表达的影响同ROS爆发具有相似的结果。说明SA可能通过调控植株抗病相关酶的基因表达及ROS爆发进而提高植株抗灰霉病能力的，而Ca~(2+)通过促进SA诱导植株抗病相关酶基因表达及ROS爆发而进一步提高植株抗灰霉病能力。
5.明确了Ca~(2+)对SA诱导番茄抗灰霉病蛋白表达的调控作用。采用TCA/丙酮法提取各处理接种灰霉菌48h后番茄幼苗叶片的蛋白，并采用双向电泳技术及PD-Quest软件分析，发现每张图谱可检测到超过500个蛋白点，软件分析得出65个差异蛋白点，在两个处理中均表达上调的蛋白有19个；两个处理都表达下降的蛋白点有6个；两个处理均是新出现的蛋白点有9个；此外，在SA处理中，有2个蛋白点仅在该处理中呈上调表达，有3个蛋白点仅在该处理中呈下调表达，有2个蛋白点仅在该处理中是新出现的蛋白点；有2个蛋白点在该处理中是消失的蛋白点。在Ca+SA处理中，有8个蛋白点仅在该处理中呈上调表达，有1个蛋白点在该处理中呈下调表达，10个蛋白点仅在Ca+SA处理中是新出现的蛋白点，有3个蛋白点仅在该处理中表达消失。
6.采用基质辅助激光解析电离飞行时间质谱技术(MODI-TOF-TOF)将稳定表达、丰度值变化1.5倍以上的蛋白点，并根据以往经验排除功能性蛋白，挑选出其中的50个进行鉴定，根据NCBI数据库搜索比对，鉴定成功了42个蛋白质点。它们分别参与了植物自身的防卫反应、能量代谢、蛋白质合成和转录调控等多个生理生化过程。明确了病程相关蛋白5和STH2、酸性内源几丁质酶等抗病相关物质以及SOD、APX等ROS相关酶在Ca~(2+)促进SA诱导番茄抗病性中起到了关键性的作用。
7.营养液Ca~(2+)浓度处理配合施用SA对NahG及其野生型（Moneymaker）的病情指数调查表明，在SA存在条件下Ca~(2+)可显著提高番茄抗病性。分析了SA依赖的信号途径下游关键基因(NPR1、PR1、TGA1a、TGA2.2)及蛋白激酶基因（MPK2、MPK4）的表达情况。NPR1、PR1、TGA1a、TGA2.2在补Ca~(2+)处理中表达均明显上调，而MPK4则表达趋势与此相反，仅在0mM Ca~(2+)处理中表达才上调，而0mM Ca~(2+)处理番茄抗病性显著降低。证明了SA信号转导相关基因及MPK2基因表达上调是营养液Ca~(2+)浓度处理配合施用SA提高番茄抗病性的主要原因，且与MPK4无关。
8.明确了营养液Ca~(2+)浓度处理再增施SA对Ca~(2+)受体相关基因表达的影响。与3mMCa~(2+)处理相比8mM Ca~(2+)处理明显提高了CDPK的表达水平，0mM Ca~(2+)处理中则明显下降；CaM的表达显示其对外源SA处理较为敏感；CBL的上调表达可能受Ca~(2+)和SA的共同作用。因此推测营养液Ca~(2+)配合SA处理提高番茄抗灰霉病可能部分通过Ca~(2+)受体调节SA信号途径相关基因表达。
9.明确了营养液Ca~(2+)处理均可显著降低野生型‘Moneymaker’和SA缺失突变体‘NahG’品种的病情指数，但野生型‘Moneymaker’比SA缺失突变体‘NahG’的病情指数更低；Ca+SA处理进一步显著降低番茄野生型‘Moneymaker’和SA缺失突变体‘NahG’品种的病情指数，同样野生型‘Moneymaker’比SA缺失突变体‘NahG’的病情指数更低；说明Ca~(2+)既有与SA共同降低番茄病情指数的作用，Ca~(2+)也具有不依赖SA降低番茄病情指数的作用，详细机制有待进一步研究。
Tomao (Solanum lycopersicum Mill) is an important kind in horticulture production inthe world and China. However, tomato is also a frequently-occurring crop. Currently, theproducer mainly depended on the chemicals to prevent disease，which lead to environmentaland products pollution. The induced resistance is an action mode that comes from plantsthemselves with the characteristic of activity. And which was worth to be researching andapplication. We should carry on depth research on the mechanism of induced resistance if wewant to exploit the plant induce resistance in order to enhance plant defence.
Ca~(2+)and Salicylic acid are considered that the important resistance substance (Sugimotoet al.,2005; Volpin and Elad,1991; Yoon CS et al.,2010; White,1979; Malamy et al.,1990;Gaffney et al.,1993; Durrant2004; Mauch2001). A growing number of detail reports indicatethat Ca~(2+)have strength role on SA induced effects, such as ROS, sub-high temperatureresistance, drought resistance, on-off of stomata and pos-harvest physiology of peach (Pei etal.,2000；Liu et al.,2005; Wang et al.,2003; Liu et al.,2003; Ao,2003). There is not detailreport on the mechanism of Ca~(2+)effect on SA induced resistance on Botrytis cinerea infectedtomato, and research of the mechanism is lack.
The cultivated tomato ‘L402’ which was sensitive to Botrytis cinerea, was employed inthis experiment, the treatments of with or without Ca~(2+)was designed, base on the level ofphysiology and molecule analysised the Ca~(2+)effect on SA induced resistance on Botrytiscinerea inoculation. It is helpful to study the mechanism to achieve the finally objective toenhanced the SA resistance efficiency and the yield and qulity both in scientific and practicalfields. The main research results are as follows:
1. It was clear that Ca~(2+)have the ablity to enhanced plants resistance through anaylisis thedisease index of different Ca~(2+)concentration of nutrient solution and foliar application Ca~(2+).Compare with control, the disease index declined63.61%in the treatment of supplementingCa~(2+)to nutrient solution; and the disease index reduced6.04%in treatment of foliarapplication Ca~(2+)compare to control; which illustrated that it was efficiency to apply Ca~(2+)tothizosphere. It was found that Magnesium (Mg~(2+)) did negative effect on tomato resistance onBotrytis cinerea, in alone Mg~(2+)treatment the disease index was78.25，which was increase4.54%compares with control.
2. Aimed to analysis the detail reasons of effectively enhancing the tomato resistanceunder thizosphere Ca~(2+)supplementation. It was found that the effect of Ca~(2+)treatment on SAaccumulation and SA synthesis in tomato leaves. It was showed that the expression level ofCM、PAL and activity of BA2H increased significantly Ca~(2+)treated, however, the expressionlevel of PAD4and ICS1were not significantly. So SA accumulation in Ca~(2+)concentrationtreated, which was the results of PAL pathway related genes expression levels’ increase.
3. It was clear that the disease index reduced significantly by foliar appling SA; andMg~(2+)did negative effect on SA induced resistance on Botrytis cinerea. The efficiency was difference that Ca~(2+)and SA in different order were applied to test the resistance on Botrytiscinerea. The Ca+SA treatment have the best effect on defense, the disease index decrease17.93%compare with alone SA treatment, and13.34%compare with SA+Ca, there was notdifference between SA+Ca and SA treated. The Mg~(2+)declined SA-induced resistance on graymold; the results of application of SA after Ca~(2+)indicated that Ca~(2+)enhanced SA-inducedresistance on gray mold in tomato.
4. It was found that Ca~(2+), EGTA, SA alone and applied complex led to ROS, enzymesactivity and genes expression increase under induced treatments or inoculation with graymold after induced treatments. Among the different treatments, the results showed SA caninduce ROS and Ca~(2+)enhanced the SA-induced ROS. In contrast，pretreatment with Ca~(2+)chelator EGTA, weakened the ROS and weakened the SA-induced ROS. The similar resultswere found in PAL, chintase, β-1,3-glucancse activity and genes expressions (PR1、 PR2aand PR3b). It indicated that SA enhanced tomato resistance by regulation of ROS andpathogensis related protein activity and expression, and Ca~(2+)promoted SA-induced resistanceto tomato gray mold by further increase these index.
5. It was clear that the function of proteins expression of Ca~(2+)regulating SA-inducedresistance to gray mold. The tomato leaves protein was extracted with TCA/acetone under theCK、SA、Ca+SA treatments inoculation with gray mold after48h. There were more than500protein spots in gel image through2-DE technical and the software of PD-Quest. We totallyfound out65differentially expressed protein spots. In detail, there were19spots wup-regulated,6spots down-regulated,9new spots in SA and Ca+SA treatemts; in addition,2spots up-regulated,3spots down-regulated,2new spots and2spots disappeared only in SAtreatment; and8spots up-regulated,1spots down-regulated,10new spots and3spotsdisappeared only in Ca+SA treatemts.
6.50differentially expressed protein spots with stable expression and higher than1.5-fold spot density were analysed by MODI-TOF-TOF. The42protein spots were obtainedby NCBI database searehing and intereomparison preliminary identifieation. These proteinswere associated with a variety of functions, including photosynthesis, energy metabolism,metabolism, ROS, protein synthesis and degradation, cell growth/division, signal transduction,and transcriptional. The key protein and enzyme of Ca~(2+)enhanced SA-induced resistance ontomato Botrytis cinerea were pathogenesis-related protein5, PR STH-2, acidic endochitinase,SOD and APX.
8. It was found that the disease index of treatments of Ca~(2+)concentration of nutrientsolution with SA application reduced significantly in ‘NahG’ and WT, which proved that Ca~(2+)enhanced tomato resistance at the condition of SA presence. Effects of Ca~(2+)on SA signalingtransduction in tomato was observed in this paper. The SA-dependent signaling related keygene (NPR1、PR1、TGA1a、TGA2.2) and two mitogen-activated protein kinase (MPK2、MPK4)expression level were analysised. An obvious increase of NPR1、PR1、TGA1a、TGA2.2、MPK2transcript was observed in8mM Ca~(2+)treated tomato seedlings. In contrary, the MPK4 expression level was up-regulation only in0mM Ca~(2+)treated, and the resistance declinedsignificantly in the treatment of0mM Ca~(2+). Looking From the rule of changes, Ca~(2+)concentration of nutrient solution with SA application enhanced tomato resistance, which wasresults of up-regulation expression of SA signaling pathway related genes and MPK2, not theMPK4.
9. The gene expression of Ca~(2+)related receptor had a great change after treatment ofCa~(2+)concentration of nutrient solution with SA application. The transcription of CDPKenhanced obviously in8mM Ca~(2+)comparation with3mM Ca~(2+), which expression level waslower in0mM Ca~(2+)than3mM Ca~(2+)treatment. From the analysis of relationship betweenCa~(2+)concentration and CaM expression level, it was indicated that the CaM transcription waspositively correlated to SA simulation. The most probably of CBL transcription situation wasCa~(2+)and SA co-regulation. So we speculated that Ca~(2+)related receptor partly participated inthe process of Ca~(2+)concentration with SA application treatment to enhance tomato resistance.
7. It was clear that Ca~(2+)have the ablity to enhanced plants resistance through treated‘Moneymaker’ and’NahG’ with different Ca~(2+)concentration of nutrient solution andanaylisis the disease index. The disease was lower in ‘Moneymaker’comparation to ‘NahG’.Further more, the disease index was lower in ‘Moneymaker’ and’NahG’ treated with2mMSA after appling with different Ca~(2+)concentration of nutrient solution comparation totreatments of different Ca~(2+)concentration of nutrient solution, it showed that Ca~(2+)not onlyhad the ablity to reduce tomato disease index combined with SA, but also had the function toinclined tomato disease index by the mode of SA-independent. The detail mechanism need tofurther research.
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