石蒜花发育生理及切花栽培关键技术研究
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摘要
石蒜(Lycoris radiata)花葶粗壮,花型优美,花色艳丽,抗逆性强,且具较高的药用价值,开发潜力巨大,但其切花栽培研究严重滞后。本项目以四年生石蒜鳞茎为研究对象,从发育物候学、形态解剖、发育生理、贮藏生理、切花栽培等方面对石蒜花发育生理进行了比较系统的研究,初步探讨了石蒜的花发育机理及切花栽培技术,以期为石蒜的开发利用和花发育机理研究提供资料参考。
主要结论如下：
1.石蒜的物候观察与结构解剖研究表明,石蒜具有适应阴湿环境的叶片结构特征。石蒜整个生育期包括花芽分化期(3月下旬～5月中旬)、花器官发育期(5～7月上旬)、抽苔期(8月中、下旬)、开花期(9月上旬～10月上旬)、出叶期(10月中、下旬)、绿叶期(11上旬～3月中旬)、叶枯黄期(3月下旬～5月上旬)。石蒜的生长发育受环境温度的影响较大。当月日均温度分别为18.3℃～21.8℃、10.6℃～14.8℃、5℃～9℃、22.1℃～25℃、25℃～31℃,石蒜叶片生长分别处于出叶期、旺盛生长期、缓慢生长期、叶黄期和叶枯萎期；而在花原体形成之前需要12.88℃以下的低温刺激,雄蕊形成后则需要17.3℃～23.35℃的适当升温,而在抽葶及开花期又需要25℃～30℃的高温环境。此外,持续28℃～30℃左右的月日均极端气温及1.69℃左右的土—气间的温差也可能是生长点转相的拐点。
2.石蒜发育生理研究表明,鳞茎内还原性糖与可溶性蛋白含量,SOD与POD活性及激素含量的变化在一定程度上标志了石蒜发育的不同阶段。其中,在花芽分化期,还原性糖与可溶性蛋白质含量均分别呈逐渐上升趋势,SOD和POD活性则相反；而在开花期至出叶期,前者平稳下降,后者则呈明显上升趋势。发育期间鳞茎芽内激素的峰值依序出现(IAA→GA→ZR→ABA→IPA),且IAA与ABA、ZR负相关,而ABA与IPA、GA正相关。具体表现为相对高比值的IAA/GA、IAA/ZR、GA/IPA有利于石蒜花芽的形态分化,而高比值的ABA/GA、ZR/IPA、ABA/ZR则有利于花形态建成、开花及花后萌叶。
3.鳞茎贮藏试验表明,温度和贮藏时间均会明显影响石蒜的花期与切花品质。5℃的低温贮藏虽可明显推迟花期和展叶期,却会相应降低花葶及叶片的长度；14℃贮藏处理虽可提前萌叶(+29d),但当年不开花；而22℃～26℃的高温贮藏则可提前10～16d开花,且切花品质也会随贮藏时间的延长而下降(如开花率低、花期短)。
4.贮藏生理研究表明,鳞茎内可溶性糖、可溶性蛋白质、POD、SOD活性、MDA含量均表现于随着贮藏温度的升高及时间的延长其含量或活性先升后降的趋势,但峰值出现的温度值不同。IAA/ZR、ZR/IPA、ABA/GA在石蒜鳞茎休眠解除的过程中具有重要作用。在5℃～14℃的低温范围内,鳞茎芽内IAA、ABA、IPA含量及ABA/GA随贮藏温度的升高、时间的延长而增加,而GA和ZR却相反；且IAA、ABA与ZR含量负相关。
5.切花栽培试验表明,遮光、施肥、施肥×遮光三因子均可有效地促进石蒜叶片生长、花期延迟、开花整齐及切花品质的提高,其效应表现于遮光>施肥>施肥*遮光。四种遮光处理中,以遮光45%的处理效应最佳,不仅花期显著推迟,抽葶率最高(达66.76%),且切花品质最好、鳞茎产量与品质也最高,其根系生长也最好。12种施肥处理中,以B3(有机肥2.6Kg)处理的鳞茎品质最好,鳞茎数也最多；而切花品质以B12(有机肥2.6Kg+N11.7g+P10.76g+K35.8g)处理最佳,其平均花葶最长、单葶小花数也最多,花径宽度、且花葶基粗和颈粗也显著大于对照。在所有处理中,以A2B3(遮光45%+有机肥2.6Kg)处理的切花产量最高(达104.43支·m-2)和鳞茎品质最好,花葶也较粗,其根系数量也最多；而以A2B12(遮光%+有机肥2.6Kg+N11.7g+P10.76g+K35.8g)的切花品质最高。综合考虑以A2B3处理效果最优。
Lycoris radiata, which has stout flower stems, gorgeous color, high medical value and strong resistance, is considered to be of great development potential. But the research on cut-flower cultivation technology is seriously lagged behind. This paper takes four-year-old asexual bulbs of L. radiata as the subject and studies its floral development physiology and cut-flower cultivation techniques by examining its growth phenology, anatomical structure, development physiology, storage physiology, and cut-flower cultivation techniques. The results are supposed to provide some reference for the exploitation and flower development study of L. radiata.
The main contents and results are as follows:
1. Phenological observations and the anatomical structure show that such structural features as thinner palisade tissue, osteoporosis sponge tissue and the convex shape feature of epidermal cells explains its capacity to adapt to the damp environment. The whole growth and development period of L. radiata includes the following stages:the period of flower bud differentiation(from late March to mid May), floral organ development period(from late May to early July), bolting period(from mid to late August), blossoming stage (from early September to early October), leaf sprouting stage (from mid to late October)and green leaf stage (from November to March), and leaf scorch stage((from late March to early May). In addition, the growth and development of L.radiata are greatly affected by the environmental temperature.It's is at the respectively sprouting-leaf atage, vigorous growth stage, the slow growth stage, leaf yellowing stage, and leaf blight stage when the temperature ranges respectively from18.3℃to21.8℃, from10.6℃to14.8℃from5℃to9℃, from22.1℃to25℃, and from25℃to31℃. On the other hand, when it is respectively in the period of flower primordium, perianth form, and floral organ development, only when the temperature is also respectively below12.88℃,17.3℃～23.35℃, and25℃～30℃. In addition, the continuous high temperature or the temperature difference of1.69℃between the soil and air is probably necessary for its lower bud differentiation.
2. Studies on developmental physiology of L. radiata's bulbs show that the content of soluble sugar and soluble protein in bulbs changes when it's at different developmental stages as well as the activities of SOD and POD. In which, the content of soluble sugar and soluble protein both tend to increase gradually at the stage of flower bud differentiation to maintain relatively stable in floral development period, and to decrease significantly in the green period. But, the activities of SOD and POD decrease during flower bud differentiation period and they increase gradually during the floral development to green period. Studies of endogenous hormones content in the process of bulb development show that the peak sequence of five kinds of endogenous hormones content appears orderly as following:IAA→GA→ZR→ABA→IPA. There is a negative correlation between IAA, ABA, and ZR, and there is a positive correlation between ABA, IPA and GA content in bulb bud. The result shows the relatively high ratio of IAA/GA, IAA/ZR, GA/IPA content and ABA/GA, ZR/iPA accelerate flower bud differentiation, and high ABA/GA, ZR/IPA, ABA/ZR accelerate flower bud development, blossom and leaf sprouting.
3. The test of bulb storage shows that temperature and storage time both have an obvious influence on the florescence and quality of cut-flower of Lycoris radiata. Although the treatment of5℃cold storage significantly delays its florescence and leaf sprouting period. At the same time it also reduces accordingly the length of scape and leaf. The processing of14℃helps sprout leaves ahead of29days, but not blossom before spring. However, The high temperature (22℃～26℃) storage blooms ahead of10to16days. In addition, the quality of cut-flowers decreases with a longer storage time under the same storage temperature(the lower blossom rate, the shorter florescence).
4. Bulb storage physiological studies suggest that the change trends about the soluble sugar content, soluble protein content, POD activity, SOD activity, and MDA content during storage are similar. That is, they decrease with the increasing temperature and time. But the temperatures of their peak are different.On the other hand, the IAA/ZR, ZR/iPA and AB A/G A play an important role in the process of breaking dormancy. At low temperature of5℃to14℃, the contents of IAA, ABA, and IPA increase with temperature and so does storage time. and the change trend about ABA/GA is similar. While, the content and the with temperature and storage time increase, while the change trends of GA and ZR are the opposite. The contents of IAA and ABA are respectively negative correlation to that of ZR.
5. The experiment of cut-flower cultivation shows that three factors can be effective in promoting the growth of leaves, flowering delay, flowering uniformity, the quality of cut-flowers.The performance of effect by the strength is as the following: shading>fertilization> shading*fertilization. In four kinds of shading, the shading of45%has the best effect comprehensively. Because it not only has the highest rate of blossom(66.76%), the highest number of bulbs (67.75), and the biggist fresh weight(1184.5g), the root number and the length of roots are also the biggest, and the quality of cut-flowers is the best. For example, compared to the control, its average scape length, flower diameter, basic coarse, carotid diameter and the number of floret are respectively more6.31cm,0.3cm,0.03cm,0.02cm,0.13).In12fertilization treatments, the effect of B3on the quality of bulbs is the best (the average diameter is largest,3.55cm). Its total number of bulbs is biggest (61.5particles) and the flower number per scape is most. However, the quality of cut-flower of B12(Shading45%+Organic manure2.6kg·plt-1+N11.7g+P10.76g+K35.8g) is the best. Its average length, the flower number of every flower sequence, the flower diameter, the basic crude and the neck roughness of scape are significantly greater than that of the control.In all treatments, the processing of A2B3(Shading45%+Organic manure2.6kg·plot-1) has the biggest production of cut-flower(104.43branch·m-2), the maximum diameter of bulb, the largest number of root, and coarse scape. However, the processing of A2B12(Shading45%+Organic manure2.6kg·plot-1+N11.7g+P10.76g+K35.8g) has the superior quality of cut-flower(such as the longest scape, the flower number per scape), the biggest leaf area and bulb weight. Overall, the promoting effect of A2B12is best.
In conclusion, Lycoris radiata has some characteristics in leaf structure of adapting to the damp environment, and it nourishes with humus-rich sandy loam. Therefore, during its cut-flower cultivation, the moderate shade (45%), chemical fertilizer application amount of organic fertilizer (3.25kg·m-2) is the effectively way to cultivate the excellent quality and high yield cut-flowers, as well as delaying flowering and improving its blossom uniformity. In addition, the development of flower and leaf is mainly controlled by the balance of hormones ratio, and the requirement on the temperature in different stage of the flower bud differentiation and development is not same, the process is significantly affected by the environmental temperature. The specific remains to be further research.
主要结论如下：
1.石蒜的物候观察与结构解剖研究表明,石蒜具有适应阴湿环境的叶片结构特征。石蒜整个生育期包括花芽分化期(3月下旬～5月中旬)、花器官发育期(5～7月上旬)、抽苔期(8月中、下旬)、开花期(9月上旬～10月上旬)、出叶期(10月中、下旬)、绿叶期(11上旬～3月中旬)、叶枯黄期(3月下旬～5月上旬)。石蒜的生长发育受环境温度的影响较大。当月日均温度分别为18.3℃～21.8℃、10.6℃～14.8℃、5℃～9℃、22.1℃～25℃、25℃～31℃,石蒜叶片生长分别处于出叶期、旺盛生长期、缓慢生长期、叶黄期和叶枯萎期；而在花原体形成之前需要12.88℃以下的低温刺激,雄蕊形成后则需要17.3℃～23.35℃的适当升温,而在抽葶及开花期又需要25℃～30℃的高温环境。此外,持续28℃～30℃左右的月日均极端气温及1.69℃左右的土—气间的温差也可能是生长点转相的拐点。
2.石蒜发育生理研究表明,鳞茎内还原性糖与可溶性蛋白含量,SOD与POD活性及激素含量的变化在一定程度上标志了石蒜发育的不同阶段。其中,在花芽分化期,还原性糖与可溶性蛋白质含量均分别呈逐渐上升趋势,SOD和POD活性则相反；而在开花期至出叶期,前者平稳下降,后者则呈明显上升趋势。发育期间鳞茎芽内激素的峰值依序出现(IAA→GA→ZR→ABA→IPA),且IAA与ABA、ZR负相关,而ABA与IPA、GA正相关。具体表现为相对高比值的IAA/GA、IAA/ZR、GA/IPA有利于石蒜花芽的形态分化,而高比值的ABA/GA、ZR/IPA、ABA/ZR则有利于花形态建成、开花及花后萌叶。
3.鳞茎贮藏试验表明,温度和贮藏时间均会明显影响石蒜的花期与切花品质。5℃的低温贮藏虽可明显推迟花期和展叶期,却会相应降低花葶及叶片的长度；14℃贮藏处理虽可提前萌叶(+29d),但当年不开花；而22℃～26℃的高温贮藏则可提前10～16d开花,且切花品质也会随贮藏时间的延长而下降(如开花率低、花期短)。
4.贮藏生理研究表明,鳞茎内可溶性糖、可溶性蛋白质、POD、SOD活性、MDA含量均表现于随着贮藏温度的升高及时间的延长其含量或活性先升后降的趋势,但峰值出现的温度值不同。IAA/ZR、ZR/IPA、ABA/GA在石蒜鳞茎休眠解除的过程中具有重要作用。在5℃～14℃的低温范围内,鳞茎芽内IAA、ABA、IPA含量及ABA/GA随贮藏温度的升高、时间的延长而增加,而GA和ZR却相反；且IAA、ABA与ZR含量负相关。
5.切花栽培试验表明,遮光、施肥、施肥×遮光三因子均可有效地促进石蒜叶片生长、花期延迟、开花整齐及切花品质的提高,其效应表现于遮光>施肥>施肥*遮光。四种遮光处理中,以遮光45%的处理效应最佳,不仅花期显著推迟,抽葶率最高(达66.76%),且切花品质最好、鳞茎产量与品质也最高,其根系生长也最好。12种施肥处理中,以B3(有机肥2.6Kg)处理的鳞茎品质最好,鳞茎数也最多；而切花品质以B12(有机肥2.6Kg+N11.7g+P10.76g+K35.8g)处理最佳,其平均花葶最长、单葶小花数也最多,花径宽度、且花葶基粗和颈粗也显著大于对照。在所有处理中,以A2B3(遮光45%+有机肥2.6Kg)处理的切花产量最高(达104.43支·m-2)和鳞茎品质最好,花葶也较粗,其根系数量也最多；而以A2B12(遮光%+有机肥2.6Kg+N11.7g+P10.76g+K35.8g)的切花品质最高。综合考虑以A2B3处理效果最优。
Lycoris radiata, which has stout flower stems, gorgeous color, high medical value and strong resistance, is considered to be of great development potential. But the research on cut-flower cultivation technology is seriously lagged behind. This paper takes four-year-old asexual bulbs of L. radiata as the subject and studies its floral development physiology and cut-flower cultivation techniques by examining its growth phenology, anatomical structure, development physiology, storage physiology, and cut-flower cultivation techniques. The results are supposed to provide some reference for the exploitation and flower development study of L. radiata.
The main contents and results are as follows:
1. Phenological observations and the anatomical structure show that such structural features as thinner palisade tissue, osteoporosis sponge tissue and the convex shape feature of epidermal cells explains its capacity to adapt to the damp environment. The whole growth and development period of L. radiata includes the following stages:the period of flower bud differentiation(from late March to mid May), floral organ development period(from late May to early July), bolting period(from mid to late August), blossoming stage (from early September to early October), leaf sprouting stage (from mid to late October)and green leaf stage (from November to March), and leaf scorch stage((from late March to early May). In addition, the growth and development of L.radiata are greatly affected by the environmental temperature.It's is at the respectively sprouting-leaf atage, vigorous growth stage, the slow growth stage, leaf yellowing stage, and leaf blight stage when the temperature ranges respectively from18.3℃to21.8℃, from10.6℃to14.8℃from5℃to9℃, from22.1℃to25℃, and from25℃to31℃. On the other hand, when it is respectively in the period of flower primordium, perianth form, and floral organ development, only when the temperature is also respectively below12.88℃,17.3℃～23.35℃, and25℃～30℃. In addition, the continuous high temperature or the temperature difference of1.69℃between the soil and air is probably necessary for its lower bud differentiation.
2. Studies on developmental physiology of L. radiata's bulbs show that the content of soluble sugar and soluble protein in bulbs changes when it's at different developmental stages as well as the activities of SOD and POD. In which, the content of soluble sugar and soluble protein both tend to increase gradually at the stage of flower bud differentiation to maintain relatively stable in floral development period, and to decrease significantly in the green period. But, the activities of SOD and POD decrease during flower bud differentiation period and they increase gradually during the floral development to green period. Studies of endogenous hormones content in the process of bulb development show that the peak sequence of five kinds of endogenous hormones content appears orderly as following:IAA→GA→ZR→ABA→IPA. There is a negative correlation between IAA, ABA, and ZR, and there is a positive correlation between ABA, IPA and GA content in bulb bud. The result shows the relatively high ratio of IAA/GA, IAA/ZR, GA/IPA content and ABA/GA, ZR/iPA accelerate flower bud differentiation, and high ABA/GA, ZR/IPA, ABA/ZR accelerate flower bud development, blossom and leaf sprouting.
3. The test of bulb storage shows that temperature and storage time both have an obvious influence on the florescence and quality of cut-flower of Lycoris radiata. Although the treatment of5℃cold storage significantly delays its florescence and leaf sprouting period. At the same time it also reduces accordingly the length of scape and leaf. The processing of14℃helps sprout leaves ahead of29days, but not blossom before spring. However, The high temperature (22℃～26℃) storage blooms ahead of10to16days. In addition, the quality of cut-flowers decreases with a longer storage time under the same storage temperature(the lower blossom rate, the shorter florescence).
4. Bulb storage physiological studies suggest that the change trends about the soluble sugar content, soluble protein content, POD activity, SOD activity, and MDA content during storage are similar. That is, they decrease with the increasing temperature and time. But the temperatures of their peak are different.On the other hand, the IAA/ZR, ZR/iPA and AB A/G A play an important role in the process of breaking dormancy. At low temperature of5℃to14℃, the contents of IAA, ABA, and IPA increase with temperature and so does storage time. and the change trend about ABA/GA is similar. While, the content and the with temperature and storage time increase, while the change trends of GA and ZR are the opposite. The contents of IAA and ABA are respectively negative correlation to that of ZR.
5. The experiment of cut-flower cultivation shows that three factors can be effective in promoting the growth of leaves, flowering delay, flowering uniformity, the quality of cut-flowers.The performance of effect by the strength is as the following: shading>fertilization> shading*fertilization. In four kinds of shading, the shading of45%has the best effect comprehensively. Because it not only has the highest rate of blossom(66.76%), the highest number of bulbs (67.75), and the biggist fresh weight(1184.5g), the root number and the length of roots are also the biggest, and the quality of cut-flowers is the best. For example, compared to the control, its average scape length, flower diameter, basic coarse, carotid diameter and the number of floret are respectively more6.31cm,0.3cm,0.03cm,0.02cm,0.13).In12fertilization treatments, the effect of B3on the quality of bulbs is the best (the average diameter is largest,3.55cm). Its total number of bulbs is biggest (61.5particles) and the flower number per scape is most. However, the quality of cut-flower of B12(Shading45%+Organic manure2.6kg·plt-1+N11.7g+P10.76g+K35.8g) is the best. Its average length, the flower number of every flower sequence, the flower diameter, the basic crude and the neck roughness of scape are significantly greater than that of the control.In all treatments, the processing of A2B3(Shading45%+Organic manure2.6kg·plot-1) has the biggest production of cut-flower(104.43branch·m-2), the maximum diameter of bulb, the largest number of root, and coarse scape. However, the processing of A2B12(Shading45%+Organic manure2.6kg·plot-1+N11.7g+P10.76g+K35.8g) has the superior quality of cut-flower(such as the longest scape, the flower number per scape), the biggest leaf area and bulb weight. Overall, the promoting effect of A2B12is best.
In conclusion, Lycoris radiata has some characteristics in leaf structure of adapting to the damp environment, and it nourishes with humus-rich sandy loam. Therefore, during its cut-flower cultivation, the moderate shade (45%), chemical fertilizer application amount of organic fertilizer (3.25kg·m-2) is the effectively way to cultivate the excellent quality and high yield cut-flowers, as well as delaying flowering and improving its blossom uniformity. In addition, the development of flower and leaf is mainly controlled by the balance of hormones ratio, and the requirement on the temperature in different stage of the flower bud differentiation and development is not same, the process is significantly affected by the environmental temperature. The specific remains to be further research.
引文
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