根域限制对牡丹碳代谢和内源激素变化的影响
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摘要
以‘洛阳红’牡丹为材料，研究了根域限制（盆栽）对牡丹碳代谢和内源激素变化的影响。结果表明，根域限制使牡丹的株高和冠幅、混合芽的生长发育以及叶面积大小都受到抑制。一二级根的数量减少，一级根的干重和鲜重显著低于对照，二级根的干重和鲜重与对照无显著差异；三级根的数量增加，干重显著低于对照。一级根和三级根的含水量均高于对照，二级根的含水量低于对照，说明牡丹干物质积累主要集中在二级根，一级根的存储干物质的能力下降，功能退化，三级根的吸水能力增加，说明根域限制并未使牡丹根部受到水分胁迫。根域限制使牡丹基质的pH值增加。
根域限制显著地降低了牡丹叶片的净光合速率（Pn），蔗糖磷酸合成酶（SPS）和酸性转化酶（AI）的活性。根域限制下，牡丹叶片中SPS和AI活性的降低，表明其源强和库强都受到抑制。盛花期之前，牡丹叶片中蔗糖、淀粉的含量显著低于对照（大田栽培）；从盛花期到开花后期，库强拉动蔗糖运输到库器官的动力不足，淀粉水解少，蔗糖、淀粉在叶片中形成累积；盛花前后己糖变化平稳，含量总体低于对照。库强小是牡丹Pn降低的主要原因，还与叶面积小和碳水化合物在叶片中的累积有一定的相关性。
根域限制下，牡丹花朵的直径和高度都显著降低，牡丹在初开期瓣干重占总干重比例显著低于对照，从圆桃期到初开期花瓣干重的增加速度也低于对照。这说明根域限制抑制了牡丹花瓣干重的增加和干物质积累的速度。从圆桃期到盛开期，雄蕊干重/总干重的比值高于对照，雌蕊干重/总干重的比值低于对照，表明根域限制下，牡丹花器官干重的分配发生了变化，干重分配到花瓣和雌蕊的比例有所降低，分配到雄蕊的比例增加。根域限制使牡丹花器官总体的蔗糖、己糖含量降低，但是降低的程度不一样，在花瓣和雌蕊中降低的程度要低于雄蕊中降低的程度。牡丹花瓣中淀粉含量在圆桃期和初开期，显著低于对照，雄蕊中淀粉含量在圆桃期、初开期和盛开期显著低于对照，雌蕊中淀粉含量在四个时期都显著低于对照。雌蕊中可溶性糖和淀粉的不足，是根域限制牡丹败育率较高的原因。从圆桃期到衰老期，根域限制牡丹花瓣、雄蕊、雌蕊中AI活性都显著低于对照，AI活性的降低使牡丹花瓣、花粉和子房的生长和发育活力都受到影响，说明根域限制牡丹花器官库强小，拉动蔗糖输送到花器官的动力不足，蔗糖输送的少，蔗糖被利用后，能形成淀粉被存储起来作为储备资源也少，碳水化合物储备资源受限，调动碳水化合物利用的动力不足，以牡丹花径小，开花不良，败育率较高等形式表现出来。
根域限制下，牡丹根中ZR、IAA和GA含量总体低于对照，其中IAA含量在四个时期与对照差异显著，下降相对较为明显；牡丹根中ABA含量总体高于对照，旺盛生长Ⅱ期和开花前期显著高于对照。牡丹茎中ZR、IAA和GA含量总体低于对照，ZR含量在萌芽期、IAA含量在在旺盛生长Ⅱ期和开花前期、GA含量在开花前期与对照差异显著；茎中ABA含量总体高于对照，在萌芽期和旺盛生长Ⅱ期与对照差异显著。叶片中IAA和GA含量略微低于对照，但与对照差异不显著；ZR含量在旺盛生长期与对照差异显著；ABA含量在旺盛生长期和开花前期显著高于对照。总体看来，根域限制降低了ZR、IAA和GA的含量（根中表现较为明显）。而增加了ABA的含量（叶片中表现较为明显）。
This study analyzed the effect of root restriction (potted cultivated) on carbohydratemetabolism and endogenous hormones in 'Luoyang hong' tree peony (Paeonia suffruticosa).Theresult showed that root restriction reduced the plant height, canopy size, growth and developmentlevel of bud of tree peony. The number of the rootⅠ,rootⅡ of tree peony were reduced. Freshand dry weight of the rootⅠ were significantly lower than control. Fresh and dry weight of theroot Ⅱ were not significantly different from the control. The number of root Ⅲ was increased,but their dry weight was significantly lower than the control. Water content of the rootⅠ and rootⅢ were higher than control. Water content of rootⅡ was lower than the control. So it can be seenthat dry matter of root restriction tree peony were mainly concentrated in the root Ⅱ, and thestorage ability of rootⅠwas decreased. The water absorption capacity of root Ⅲ was increasedwhich means that root were not suffered water stress when grown under root restriction. The pHvalue of peony matrix was increased when grown under root restriction.
Root restriction significantly reduced the net photosynthetic rate (Pn), sucrose phosphatesynthase (SPS) activity and acid invertase (AI) activity of tree peony leaves. The declined SPSactivity and AI activity in tree peony leaves show that both source and sink strength are inhibitedby root restriction. Before anthesis, sucrose and starch levels were significantly lower in rootrestriction tree peony leaves than that in control. However, from flowering-stage to late-floweringstage, sucrose and starch were accumulated in root restriction tree peony leaves mainly because ofthe low sink strength which limited the driving force to transport sucrose to sink organ and tohydrolyze starch. Hexose content in root restriction tree peony leaves were lower than the controloverall and the chagne was smooth at anthesis. Lower level of sink strength was the main reasonfor the reduced Pn of tree peony grown under root restriction. The reduction of leaf area andaccumulation of carbohydrates in leaves can also explain the decline of Pn to a certain extent.
Under root restriction, the diameter and thickness of tree peony flower were significantlyreduced. At initiating-bloom-stage, the ratio of petal dry weight to total flower dry weight wassignificantly lower than control. From round-peach stage to initiating-bloom-stage, dry matterincrease rate of petal was lower than the control, which means that root restriction inhibited drymatter increase and dry matter increase rate of petal. From round-peach stage to full-bloom stage,the ratio of stamen dry matter to total flower dry weight was higher than the control, the ratio ofpistil dry weight to total flower dry weight was lower than the control, it shows that dry matter distribution of flower organ was changed, which distributed relatively more to stamens anddistributed relatively less to petals and pistils. Sucrose content and hexose content of tree peonyflower organ were decreased. However, the reduction degrees between different flower organswere different. The reduction degree in petals and pistils were lower than the reduction degree instamens. From round-peach stage to initiating-bloom-stage, starch content in petal wassignificantly lower than the control. Starch content in stamen was significantly lower than thecontrol at round-peach stage, initiating-bloom-stage and full-bloom stage. Starch content in pistilat all the stage was significantly lower than the control. The lack of soluble content and starchcontent in pistil were the reason for the higher abortion rate of tree peony grown under rootrestriction.From round-peach stage to senescence stage, activities of AI in petals, stamens andpistils were significantly lower than that in the control. The growth and development of petals,pollen and ovary were all affected by the decreased AI activity. It can be seen that the sinkstrength of flower organ was not strong enough and have insufficient power to transport sucrose.There is less sucrose to form starch when sucrose were used. There are not enough carbohydrateand lack of motivation to mobilize carbohydrate to utilize, so the flower diameter was small,flowering badly and have higher abortion rate in tree peony when growing under root restriction.
ZR, IAA and GA content in the root of tree peony was generally lower than the controlgrowing under root restriction. IAA content in the root was significantly lower than the control inall the sampling stage and its reduction was obvious. ABA content is generally higher than thecontrol. At fast-growing stageⅡand before-flowing stage, ABA content in root were significantlyhigher than the control. ZR, IAA and GA content in the stems of tree peony was generally lowerthan the control growing under root restriction. ZR content in stems at germination stage wassignificantly lower than that in the control. IAA content in stems at germination stage andfast-growing stageⅡ were significantly lower than that in the control. GA content in stems atbefore-flowing stage was significantly lower than that in the control. IAA content and GA contentin tree peony leaves were slightly lower than that in the control, but the difference were notsignificant. ZR content in tree peony leaves at fast-growing stage was significantly lower than thatin the control. ABA content in tree peony leaves at fast-growing stage and before-flowing stagewere significantly higher than that in the control. Generally root restriction reduced ZR content,IAA content and GA content in tree peony, which is more obviously in root. ABA content weregenerally increased in tree peony when grown under restriction, which is more obviously inleaves.
根域限制显著地降低了牡丹叶片的净光合速率（Pn），蔗糖磷酸合成酶（SPS）和酸性转化酶（AI）的活性。根域限制下，牡丹叶片中SPS和AI活性的降低，表明其源强和库强都受到抑制。盛花期之前，牡丹叶片中蔗糖、淀粉的含量显著低于对照（大田栽培）；从盛花期到开花后期，库强拉动蔗糖运输到库器官的动力不足，淀粉水解少，蔗糖、淀粉在叶片中形成累积；盛花前后己糖变化平稳，含量总体低于对照。库强小是牡丹Pn降低的主要原因，还与叶面积小和碳水化合物在叶片中的累积有一定的相关性。
根域限制下，牡丹花朵的直径和高度都显著降低，牡丹在初开期瓣干重占总干重比例显著低于对照，从圆桃期到初开期花瓣干重的增加速度也低于对照。这说明根域限制抑制了牡丹花瓣干重的增加和干物质积累的速度。从圆桃期到盛开期，雄蕊干重/总干重的比值高于对照，雌蕊干重/总干重的比值低于对照，表明根域限制下，牡丹花器官干重的分配发生了变化，干重分配到花瓣和雌蕊的比例有所降低，分配到雄蕊的比例增加。根域限制使牡丹花器官总体的蔗糖、己糖含量降低，但是降低的程度不一样，在花瓣和雌蕊中降低的程度要低于雄蕊中降低的程度。牡丹花瓣中淀粉含量在圆桃期和初开期，显著低于对照，雄蕊中淀粉含量在圆桃期、初开期和盛开期显著低于对照，雌蕊中淀粉含量在四个时期都显著低于对照。雌蕊中可溶性糖和淀粉的不足，是根域限制牡丹败育率较高的原因。从圆桃期到衰老期，根域限制牡丹花瓣、雄蕊、雌蕊中AI活性都显著低于对照，AI活性的降低使牡丹花瓣、花粉和子房的生长和发育活力都受到影响，说明根域限制牡丹花器官库强小，拉动蔗糖输送到花器官的动力不足，蔗糖输送的少，蔗糖被利用后，能形成淀粉被存储起来作为储备资源也少，碳水化合物储备资源受限，调动碳水化合物利用的动力不足，以牡丹花径小，开花不良，败育率较高等形式表现出来。
根域限制下，牡丹根中ZR、IAA和GA含量总体低于对照，其中IAA含量在四个时期与对照差异显著，下降相对较为明显；牡丹根中ABA含量总体高于对照，旺盛生长Ⅱ期和开花前期显著高于对照。牡丹茎中ZR、IAA和GA含量总体低于对照，ZR含量在萌芽期、IAA含量在在旺盛生长Ⅱ期和开花前期、GA含量在开花前期与对照差异显著；茎中ABA含量总体高于对照，在萌芽期和旺盛生长Ⅱ期与对照差异显著。叶片中IAA和GA含量略微低于对照，但与对照差异不显著；ZR含量在旺盛生长期与对照差异显著；ABA含量在旺盛生长期和开花前期显著高于对照。总体看来，根域限制降低了ZR、IAA和GA的含量（根中表现较为明显）。而增加了ABA的含量（叶片中表现较为明显）。
This study analyzed the effect of root restriction (potted cultivated) on carbohydratemetabolism and endogenous hormones in 'Luoyang hong' tree peony (Paeonia suffruticosa).Theresult showed that root restriction reduced the plant height, canopy size, growth and developmentlevel of bud of tree peony. The number of the rootⅠ,rootⅡ of tree peony were reduced. Freshand dry weight of the rootⅠ were significantly lower than control. Fresh and dry weight of theroot Ⅱ were not significantly different from the control. The number of root Ⅲ was increased,but their dry weight was significantly lower than the control. Water content of the rootⅠ and rootⅢ were higher than control. Water content of rootⅡ was lower than the control. So it can be seenthat dry matter of root restriction tree peony were mainly concentrated in the root Ⅱ, and thestorage ability of rootⅠwas decreased. The water absorption capacity of root Ⅲ was increasedwhich means that root were not suffered water stress when grown under root restriction. The pHvalue of peony matrix was increased when grown under root restriction.
Root restriction significantly reduced the net photosynthetic rate (Pn), sucrose phosphatesynthase (SPS) activity and acid invertase (AI) activity of tree peony leaves. The declined SPSactivity and AI activity in tree peony leaves show that both source and sink strength are inhibitedby root restriction. Before anthesis, sucrose and starch levels were significantly lower in rootrestriction tree peony leaves than that in control. However, from flowering-stage to late-floweringstage, sucrose and starch were accumulated in root restriction tree peony leaves mainly because ofthe low sink strength which limited the driving force to transport sucrose to sink organ and tohydrolyze starch. Hexose content in root restriction tree peony leaves were lower than the controloverall and the chagne was smooth at anthesis. Lower level of sink strength was the main reasonfor the reduced Pn of tree peony grown under root restriction. The reduction of leaf area andaccumulation of carbohydrates in leaves can also explain the decline of Pn to a certain extent.
Under root restriction, the diameter and thickness of tree peony flower were significantlyreduced. At initiating-bloom-stage, the ratio of petal dry weight to total flower dry weight wassignificantly lower than control. From round-peach stage to initiating-bloom-stage, dry matterincrease rate of petal was lower than the control, which means that root restriction inhibited drymatter increase and dry matter increase rate of petal. From round-peach stage to full-bloom stage,the ratio of stamen dry matter to total flower dry weight was higher than the control, the ratio ofpistil dry weight to total flower dry weight was lower than the control, it shows that dry matter distribution of flower organ was changed, which distributed relatively more to stamens anddistributed relatively less to petals and pistils. Sucrose content and hexose content of tree peonyflower organ were decreased. However, the reduction degrees between different flower organswere different. The reduction degree in petals and pistils were lower than the reduction degree instamens. From round-peach stage to initiating-bloom-stage, starch content in petal wassignificantly lower than the control. Starch content in stamen was significantly lower than thecontrol at round-peach stage, initiating-bloom-stage and full-bloom stage. Starch content in pistilat all the stage was significantly lower than the control. The lack of soluble content and starchcontent in pistil were the reason for the higher abortion rate of tree peony grown under rootrestriction.From round-peach stage to senescence stage, activities of AI in petals, stamens andpistils were significantly lower than that in the control. The growth and development of petals,pollen and ovary were all affected by the decreased AI activity. It can be seen that the sinkstrength of flower organ was not strong enough and have insufficient power to transport sucrose.There is less sucrose to form starch when sucrose were used. There are not enough carbohydrateand lack of motivation to mobilize carbohydrate to utilize, so the flower diameter was small,flowering badly and have higher abortion rate in tree peony when growing under root restriction.
ZR, IAA and GA content in the root of tree peony was generally lower than the controlgrowing under root restriction. IAA content in the root was significantly lower than the control inall the sampling stage and its reduction was obvious. ABA content is generally higher than thecontrol. At fast-growing stageⅡand before-flowing stage, ABA content in root were significantlyhigher than the control. ZR, IAA and GA content in the stems of tree peony was generally lowerthan the control growing under root restriction. ZR content in stems at germination stage wassignificantly lower than that in the control. IAA content in stems at germination stage andfast-growing stageⅡ were significantly lower than that in the control. GA content in stems atbefore-flowing stage was significantly lower than that in the control. IAA content and GA contentin tree peony leaves were slightly lower than that in the control, but the difference were notsignificant. ZR content in tree peony leaves at fast-growing stage was significantly lower than thatin the control. ABA content in tree peony leaves at fast-growing stage and before-flowing stagewere significantly higher than that in the control. Generally root restriction reduced ZR content,IAA content and GA content in tree peony, which is more obviously in root. ABA content weregenerally increased in tree peony when grown under restriction, which is more obviously inleaves.
引文
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