树月季不同砧穗组合生长及生理生化反应的比较研究
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摘要
本试验以日本无刺蔷薇（Rosa multiflora var）（试验代号A）、美国多花无刺蔷薇（待定）（试验代号B）、美国M1号（R. multiflora var.’Dr.Huey’）（试验代号C）为砧木；以大花月季绯扇（Hiohgi）（试验代号a）、四季玫瑰（Rosa rugosa Thunb）(试验代号b)、妙峰山玫瑰（Rosa rugosa）（试验代号c）、丰花月季曼海姆(Schloss Mannheim)（试验代号d）为接穗，采用芽接法进行树月季嫁接培育。以自根苗为对照，通过砧木对接穗生物量、光合指标、保护酶系统、激素、可溶性蛋白、脯氨酸含量等生长、生理生化指标测定，及接穗对砧木增粗、根系生长的影响强度观测，探讨和揭示不同砧穗组合树状月季砧穗相互作用规律，为合理选择砧木和认识砧穗间相互作用机理，提供理论依据。主要结论如下：
（1）月季和玫瑰嫁接在蔷薇砧木上，可以明显提高接穗自身叶片净光合速率、提高抗氧化酶活性、增加激素含量，可以促进树体生长，加快砧木茎增粗及提高根系生长量。
（2）所有供试材料中嫁接苗的光合速率峰值及日平均光合速率均高于接穗自根苗。全天光合速率最高值在10.57μmolm·~(-2)·s~(-1)--13.51μmolm·~(-2)·s~(-1)之间；日平均光合速率在7.73μmolm·~(-2)·s~(-1)--9.16μmolm·~(-2)·s~(-1)之间，嫁接苗较接穗自根苗高14.57%--46.56%。各供试材料叶片可溶性蛋白含量差异显著但无明显规律，四季玫瑰嫁接苗及自根苗可溶性蛋白含量整体较高，在17.83mg·g~(-1)FM--26.02mg·g~(-1)FM之间，妙峰山玫瑰嫁接苗及自根苗叶片可溶性蛋白含量整体较低，在3.90mg·g~(-1)FM--7.77mg·g~(-1)FM之间；光合色素含量除d为3.81mg·g~(-1)FM，C+d为2.68mg·g~(-1)FM以外，其余供试材料表现为嫁接苗叶片光合色素含量高于接穗自根苗。各供试材料光合生理生化指标总体呈现出嫁接苗高于自根苗。
（3）嫁接苗叶片超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）活性高于自根苗，丙二醛（MDA）含量嫁接苗低于自根苗；砧木与其嫁接组合体叶片SOD活性差异不显著，供试材料中SOD活性最高的是B+b组合为857.67U·g~(-1)·min~(-1)·FW，SOD活性最低的是A+a组合为775.46U·g~(-1)·min~(-1)·FW。POD活性各供试材料差异显著，其中B+a组合POD活性最高为785U·g~(-1)·min~(-1)·FW，C+c组合POD活性最低为102.5U·g~(-1)·min~(-1)·FW，后者是前者的7.66倍；不同砧木同种接穗嫁接苗叶片CAT活性的变化趋势相似，相同砧木不同接穗品种嫁接苗之间差异显著，而同种接穗的嫁接苗与接穗自根苗之间CAT活性差异不显著，CAT活性最高的是B+a组合为303.75U·g~(-1)·min~(-1)·FW，最低的是d为170U·g~(-1)·min~(-1)·FW。嫁接苗中MDA含量最高的是A+a组合为33.98U·g~(-1)·min~(-1)·FW，其余嫁接组合叶片MDA含量均小于接穗自根苗。
（4）嫁接苗叶片脯氨酸积累量表现出高于自根苗的总趋势，且以玫瑰为接穗的嫁接苗叶片脯氨酸积累量高于以月季为接穗的嫁接苗。脯氨酸含量最低的是A+d，为9.98ug·g~(-1)·FW，最高的是C+c，为96.61ug·g~(-1)·FW，后者是前者的9.68倍。各供试材料硝酸还原酶活性（NR）没有明显的规律性，但嫁接苗叶片硝酸还原酶活性有高于自根苗叶片硝酸还原酶活性的趋势。
（5）脱落酸（ABA）和赤霉素（GA3）含量表现为嫁接苗高于自根苗。各供试材料中，ABA含量最高的是C+b组合为5.58μg·100g~(-1)·FW，最低的是a为1.08μg·100g~(-1)·FW，前者是后者的5.17倍，玫瑰嫁接苗叶片ABA含量整体高于月季嫁接苗；GA3含量最高的是B+c组合，为6.52μg·100g~(-1)·FW，最低的是a为2.25μg·100g~(-1)·FW，前者是后者的2.90倍。
（6）接穗对砧木生长具有明显的促进作用，且玫瑰种对促进砧木生长效果明显高于月季品种。嫁接苗砧木无论是地径生长量还是穗下5cm处径生长量都明显高于砧木自根苗相应指标。嫁接苗地径生长量最大的是B+c组合为12.20mm，自根苗地径生长量最小的是c为2.45mm，嫁接苗是自根苗的4.98倍；嫁接苗穗下5cm处径生长量最大的也是B+c组合，为9.58mm，而自根苗相应生长量最小的是c，为2.20mm，嫁接苗是自根苗的4.35倍。
In order to explore and reveal the interaction laws of different rootstocks and scionscombinations, the experiment of Tree Rose breeding was conducted with the method of budgrafting,in which4types of scions,including Hiohgi（test code a）、Rosa rugosa Thunb（testcode b）、Rose rugosa(test code c) and Schloss Mannheim(test code d), were grafted on3different kinds of rootstocks, such as Rosa spp.Rosoideae(test code A）, B(test code B)andRose multiflora var.’Dr.Huey’(test code C). Taking self-root seedlings as a comparison,some growth, physiological and biochemical indexes, including yield, photosynthetic indexes,enzyme system, hormone, soluble protein, proline content, etc, which reflect the effects ofrootstocks on scions, were observed. And other growth indexes of rootstocks, includig theground diameter, root volume, etc, which reflect the effect strength of scions on rootstocks,were also observed. It provides theoretical basis for selecting rootstocks appropriately andknowing the interaction mechanism between the rootstock and scion. The main conclusionswere as follows:
(1) After grafting Chinese rose and Rugosa rose on the rose, the blade net photosyntheticrate, the antioxidant enzymes activity, the amounts of hormone and the growth of scions wereimproved significantly. And the diameter growth and root volume of rootstocks were alsoincreased.
(2)The peak and average daily photosynthetic rate of grafted seedlings were higher than theself-rooted seedlings in all the experimented materials. The highest value of photosynthetic rateand average daily photosynthetic rate all day were fluctuated from10.57μmolm·~(-2)·s~(-1)to13.51μmolm·~(-2)·s~(-1)and from7.73μmolm·~(-2)·s~(-1)to9.16μmolm·~(-2)·s~(-1)respectively. The height ofgrafted seedlings was about14.57%-46.56%higher than self-rooted seedlings. And the solubleprotein content of all experimented materials were significantly different, but their rules were not obvious. For the soluble protein content, the grafted and self-rooted seedlings of Rosarugosa Thunb, which fluctuated between17.83mg g~(-1)FM and26.02mg g~(-1)FM, wererelatively high. Whereas, the grafted and self-rooted seedlings of Rose rugosa, whichfluctuated between3.90mg g~(-1)FM and7.77mg g~(-1)FM, were relatively low. In addition, thecontent of photosynthetic pigments of grafted seedlings were higher than the self-rootedseedlings in all the experimented materials excluding the Schloss Mannheim. Therefore, thephotosynthetic, physiological and biochemical indexes of grafted seedlings were generallyhigher than the self-rooted seedlings in all the experimented materials.
（3）The SOD and POD, CAT activity of grafted seedlings were higher than self-rootedseedlings, whereas, the MDA content of grafted seedlings was lower than self-rooted seedlings.The SOD activity of leaves between the rootstock and its grafted superoxide were notsignificant. Among then, the highest and the lowest value of SOD activity, which belong to thecombination of B+b and A+a, were857.67U·g~(-1)·min~(-1)·FW and775.46U·g~(-1)·min~(-1)·FWrespectively. But the POD activity of each experimented materials were significant. Thehighest value of POD activity, belonging to the combination of B+a was up to the785U g~(-1)min~(-1) FW, whereas, the lowest value, belonging to the combination of C+c, was102.5U g~(-1) min~(-1) FW. The value of former was7.66times than that of the latter. The CATactivity of the grafted seedlings with the same scions and different rootstocks were notsignificant. Whereas, the CAT activity of the grafted seedlings with the different scions and thesame rootstocks were significant. In addition, the CAT activity of the grafted and self-rootedseedlings with the same scions were not significant. The combination of B+a had the highestvalue of CAT activity, which was303.75U g~(-1) min~(-1) FW, whereas, the d had the lowest valueof CAT activity,which was about170U g~(-1)min~(-1) FW. In all grafted seedings, the highestcontent of MDA was the combination of A+a, which up to33.98U g~(-1)min~(-1) FW. For the rest ofcombinations, the content of MDA of the grafted seedlings were less than that of theself-rooted seedlings.
（4）The leaf proline content of grafted seedlings were higher than those self-rootedseedlings in general. And the proline content of grafted seedlings with Rugosa rose scion werehigher than those with the Chinese rose scion. The lowest and the highest value of prolinecontent, which belong to A+d and C+c, were9.98ug g~(-1) FW and96.61ug g~(-1) FWrespectively, And the latter was9.68times than that of the former. The nitrate reductaseactivity of all experimented materials had no apparent regularity, but nitrate reductase activityof grafted seedlings were higher than self-rooted seedlings.
（5）The contents of ABA and GA3of grafted seedlings were higher than those self-rootedseedlings. In all experiment materials, The highest content of ABA was the combination ofC+b, up to5.58μg100g~(-1) FW and the lowest was a, up to1.08μg100g~(-1) FW. The former was5.17times than that of the latter. The leaf ABA content of grafted seedlings with Rose scionwere higher than that of seedlings with the Chinese Rose scion as a whole; The highest contentof GA3was the combination of B+c, up to the6.52μg100g~(-1) FW and the lowest was a, up to2.25μg100g~(-1) FW, the former was2.90times than that of the latter.
（6）The scions had obvious promotive effect in rootstocks growth. And the Rose species ofscions were obviously better than that of Chinese rose species. Both the diameter growth andthe diameter increasing below the scion by5cm of grafted seedling were significantly higherthan those of self-rooted seedlings. The biggest diameter increment of grafted seedlings wasthe combination of B+c, up to12.20mm, However, The smallest diameter increment ofself-rooted seedlings was c, up to2.45mm. And the former was4.98times than that of thelatter. The biggest diameter increment below the scion by5cm of grafted seedlings was also thecombination of B+c, up to9.58mm, and the smallest diameter increment below the scion by5cm of self-rooted seedlings was also c, up to2.20mm. And the former was4.35times thanthat of the latter.
（1）月季和玫瑰嫁接在蔷薇砧木上，可以明显提高接穗自身叶片净光合速率、提高抗氧化酶活性、增加激素含量，可以促进树体生长，加快砧木茎增粗及提高根系生长量。
（2）所有供试材料中嫁接苗的光合速率峰值及日平均光合速率均高于接穗自根苗。全天光合速率最高值在10.57μmolm·~(-2)·s~(-1)--13.51μmolm·~(-2)·s~(-1)之间；日平均光合速率在7.73μmolm·~(-2)·s~(-1)--9.16μmolm·~(-2)·s~(-1)之间，嫁接苗较接穗自根苗高14.57%--46.56%。各供试材料叶片可溶性蛋白含量差异显著但无明显规律，四季玫瑰嫁接苗及自根苗可溶性蛋白含量整体较高，在17.83mg·g~(-1)FM--26.02mg·g~(-1)FM之间，妙峰山玫瑰嫁接苗及自根苗叶片可溶性蛋白含量整体较低，在3.90mg·g~(-1)FM--7.77mg·g~(-1)FM之间；光合色素含量除d为3.81mg·g~(-1)FM，C+d为2.68mg·g~(-1)FM以外，其余供试材料表现为嫁接苗叶片光合色素含量高于接穗自根苗。各供试材料光合生理生化指标总体呈现出嫁接苗高于自根苗。
（3）嫁接苗叶片超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）活性高于自根苗，丙二醛（MDA）含量嫁接苗低于自根苗；砧木与其嫁接组合体叶片SOD活性差异不显著，供试材料中SOD活性最高的是B+b组合为857.67U·g~(-1)·min~(-1)·FW，SOD活性最低的是A+a组合为775.46U·g~(-1)·min~(-1)·FW。POD活性各供试材料差异显著，其中B+a组合POD活性最高为785U·g~(-1)·min~(-1)·FW，C+c组合POD活性最低为102.5U·g~(-1)·min~(-1)·FW，后者是前者的7.66倍；不同砧木同种接穗嫁接苗叶片CAT活性的变化趋势相似，相同砧木不同接穗品种嫁接苗之间差异显著，而同种接穗的嫁接苗与接穗自根苗之间CAT活性差异不显著，CAT活性最高的是B+a组合为303.75U·g~(-1)·min~(-1)·FW，最低的是d为170U·g~(-1)·min~(-1)·FW。嫁接苗中MDA含量最高的是A+a组合为33.98U·g~(-1)·min~(-1)·FW，其余嫁接组合叶片MDA含量均小于接穗自根苗。
（4）嫁接苗叶片脯氨酸积累量表现出高于自根苗的总趋势，且以玫瑰为接穗的嫁接苗叶片脯氨酸积累量高于以月季为接穗的嫁接苗。脯氨酸含量最低的是A+d，为9.98ug·g~(-1)·FW，最高的是C+c，为96.61ug·g~(-1)·FW，后者是前者的9.68倍。各供试材料硝酸还原酶活性（NR）没有明显的规律性，但嫁接苗叶片硝酸还原酶活性有高于自根苗叶片硝酸还原酶活性的趋势。
（5）脱落酸（ABA）和赤霉素（GA3）含量表现为嫁接苗高于自根苗。各供试材料中，ABA含量最高的是C+b组合为5.58μg·100g~(-1)·FW，最低的是a为1.08μg·100g~(-1)·FW，前者是后者的5.17倍，玫瑰嫁接苗叶片ABA含量整体高于月季嫁接苗；GA3含量最高的是B+c组合，为6.52μg·100g~(-1)·FW，最低的是a为2.25μg·100g~(-1)·FW，前者是后者的2.90倍。
（6）接穗对砧木生长具有明显的促进作用，且玫瑰种对促进砧木生长效果明显高于月季品种。嫁接苗砧木无论是地径生长量还是穗下5cm处径生长量都明显高于砧木自根苗相应指标。嫁接苗地径生长量最大的是B+c组合为12.20mm，自根苗地径生长量最小的是c为2.45mm，嫁接苗是自根苗的4.98倍；嫁接苗穗下5cm处径生长量最大的也是B+c组合，为9.58mm，而自根苗相应生长量最小的是c，为2.20mm，嫁接苗是自根苗的4.35倍。
In order to explore and reveal the interaction laws of different rootstocks and scionscombinations, the experiment of Tree Rose breeding was conducted with the method of budgrafting,in which4types of scions,including Hiohgi（test code a）、Rosa rugosa Thunb（testcode b）、Rose rugosa(test code c) and Schloss Mannheim(test code d), were grafted on3different kinds of rootstocks, such as Rosa spp.Rosoideae(test code A）, B(test code B)andRose multiflora var.’Dr.Huey’(test code C). Taking self-root seedlings as a comparison,some growth, physiological and biochemical indexes, including yield, photosynthetic indexes,enzyme system, hormone, soluble protein, proline content, etc, which reflect the effects ofrootstocks on scions, were observed. And other growth indexes of rootstocks, includig theground diameter, root volume, etc, which reflect the effect strength of scions on rootstocks,were also observed. It provides theoretical basis for selecting rootstocks appropriately andknowing the interaction mechanism between the rootstock and scion. The main conclusionswere as follows:
(1) After grafting Chinese rose and Rugosa rose on the rose, the blade net photosyntheticrate, the antioxidant enzymes activity, the amounts of hormone and the growth of scions wereimproved significantly. And the diameter growth and root volume of rootstocks were alsoincreased.
(2)The peak and average daily photosynthetic rate of grafted seedlings were higher than theself-rooted seedlings in all the experimented materials. The highest value of photosynthetic rateand average daily photosynthetic rate all day were fluctuated from10.57μmolm·~(-2)·s~(-1)to13.51μmolm·~(-2)·s~(-1)and from7.73μmolm·~(-2)·s~(-1)to9.16μmolm·~(-2)·s~(-1)respectively. The height ofgrafted seedlings was about14.57%-46.56%higher than self-rooted seedlings. And the solubleprotein content of all experimented materials were significantly different, but their rules were not obvious. For the soluble protein content, the grafted and self-rooted seedlings of Rosarugosa Thunb, which fluctuated between17.83mg g~(-1)FM and26.02mg g~(-1)FM, wererelatively high. Whereas, the grafted and self-rooted seedlings of Rose rugosa, whichfluctuated between3.90mg g~(-1)FM and7.77mg g~(-1)FM, were relatively low. In addition, thecontent of photosynthetic pigments of grafted seedlings were higher than the self-rootedseedlings in all the experimented materials excluding the Schloss Mannheim. Therefore, thephotosynthetic, physiological and biochemical indexes of grafted seedlings were generallyhigher than the self-rooted seedlings in all the experimented materials.
（3）The SOD and POD, CAT activity of grafted seedlings were higher than self-rootedseedlings, whereas, the MDA content of grafted seedlings was lower than self-rooted seedlings.The SOD activity of leaves between the rootstock and its grafted superoxide were notsignificant. Among then, the highest and the lowest value of SOD activity, which belong to thecombination of B+b and A+a, were857.67U·g~(-1)·min~(-1)·FW and775.46U·g~(-1)·min~(-1)·FWrespectively. But the POD activity of each experimented materials were significant. Thehighest value of POD activity, belonging to the combination of B+a was up to the785U g~(-1)min~(-1) FW, whereas, the lowest value, belonging to the combination of C+c, was102.5U g~(-1) min~(-1) FW. The value of former was7.66times than that of the latter. The CATactivity of the grafted seedlings with the same scions and different rootstocks were notsignificant. Whereas, the CAT activity of the grafted seedlings with the different scions and thesame rootstocks were significant. In addition, the CAT activity of the grafted and self-rootedseedlings with the same scions were not significant. The combination of B+a had the highestvalue of CAT activity, which was303.75U g~(-1) min~(-1) FW, whereas, the d had the lowest valueof CAT activity,which was about170U g~(-1)min~(-1) FW. In all grafted seedings, the highestcontent of MDA was the combination of A+a, which up to33.98U g~(-1)min~(-1) FW. For the rest ofcombinations, the content of MDA of the grafted seedlings were less than that of theself-rooted seedlings.
（4）The leaf proline content of grafted seedlings were higher than those self-rootedseedlings in general. And the proline content of grafted seedlings with Rugosa rose scion werehigher than those with the Chinese rose scion. The lowest and the highest value of prolinecontent, which belong to A+d and C+c, were9.98ug g~(-1) FW and96.61ug g~(-1) FWrespectively, And the latter was9.68times than that of the former. The nitrate reductaseactivity of all experimented materials had no apparent regularity, but nitrate reductase activityof grafted seedlings were higher than self-rooted seedlings.
（5）The contents of ABA and GA3of grafted seedlings were higher than those self-rootedseedlings. In all experiment materials, The highest content of ABA was the combination ofC+b, up to5.58μg100g~(-1) FW and the lowest was a, up to1.08μg100g~(-1) FW. The former was5.17times than that of the latter. The leaf ABA content of grafted seedlings with Rose scionwere higher than that of seedlings with the Chinese Rose scion as a whole; The highest contentof GA3was the combination of B+c, up to the6.52μg100g~(-1) FW and the lowest was a, up to2.25μg100g~(-1) FW, the former was2.90times than that of the latter.
（6）The scions had obvious promotive effect in rootstocks growth. And the Rose species ofscions were obviously better than that of Chinese rose species. Both the diameter growth andthe diameter increasing below the scion by5cm of grafted seedling were significantly higherthan those of self-rooted seedlings. The biggest diameter increment of grafted seedlings wasthe combination of B+c, up to12.20mm, However, The smallest diameter increment ofself-rooted seedlings was c, up to2.45mm. And the former was4.98times than that of thelatter. The biggest diameter increment below the scion by5cm of grafted seedlings was also thecombination of B+c, up to9.58mm, and the smallest diameter increment below the scion by5cm of self-rooted seedlings was also c, up to2.20mm. And the former was4.35times thanthat of the latter.
引文
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