黄花蒿化感作用与友好栽培研究
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摘要
黄花蒿又名青蒿(Artemisia annua L.),为菊科—年生或越年生草本植物,是提取青蒿素的唯一原料。目前,全世界有40%左右的人口受到疟疾威胁,青蒿素是世界卫生组织(WHO)推荐治疗疟疾的首选药物。
我国黄花蒿种植面积和青蒿素产量约占全世界的90%以上,重庆市是中国黄花蒿的优质产区,种植面积和青蒿素产量分别占全国的70%和80%。青蒿素是黄花蒿(?)最多,在土壤中存在时间最长的化感物质,降低土地生产力,抑制周围和后茬植物生长,危害土壤和水体生态环境。在人工种植黄花蒿的过程中,随着青蒿素的含量和产量的提高,释放到环境中青蒿素大幅度增加,化感危害更加严重。论文采用生态学野外样方调查、培养试验、盆栽试验和田间试验等方法,研究了野生黄花蒿的群落特征及周围植物的存在状况,青蒿素对陆生和水生植物的化感效应及机理,轮作种植模式对后茬作物及自身的影响、施肥技术和水分管理对黄花蒿生长、青蒿素含量和产量,初步形成了青蒿素高产、土地利用高效的友好栽培技术。主要结果如下：
对南方9个省市48个样地野生黄花蒿种群调查发现,黄花蒿群落处于相对稳定状态。在野生黄花蒿不同群落中,出现频率最高的9个种对中,5种为菊科植物,3种禾本科植物,联结性较低。在野生黄花蒿群落中和周围,十字花科和茄科等植物极少,表明黄花蒿对其他植物具有化感效应,抑制植物的生长发育。
在土壤中添加黄花蒿产生的化感物质—青蒿素,不同程度抑制小麦、油菜和茎瘤芥等3种后茬作物的生长,生物量降幅高达41.1-51.7%,且对茎瘤芥的抑制作用低于油菜和小麦。青蒿素抑制其它植物生长,有利于扩大黄花蒿生存空间,增强竞争优势。但是,在黄花蒿种植区,选择抗化感作用较强的后茬作物可提高土地利用率和整体生产力。高浓度的青蒿素降低后茬作物叶绿素但提高类胡萝卜素含量,说明青蒿素促进叶绿素分解,导致光合速率下降,干物质积累减少。此外,青蒿素显著抑制氮、磷、钾养分吸收,与根系活力和硝酸还原酶活性降低有关。在黄花蒿收获后,适度延迟播种和移栽后茬作物可以减轻对后茬作物的危害。
在人工培养液中,不同浓度青蒿素显著抑制绿藻生长,绿藻密度可降低74.5%(蛋白核小球藻)和61.8%(斜生栅藻)。青蒿素使绿藻培养液溶解氧减少,绿藻叶绿素和蛋白质含量降低,说明青蒿素抑制绿藻生长与光合作用降低,蛋白合成受到干扰有关,大规模种植黄花蒿会降低水体自净能力和生产力,加剧污染负荷,鱼类减产和灭绝等；但分散种植和精细采收,有益于降低水体青蒿素浓度,减轻水环境风险。此外,青蒿素提高培养液电导率和绿藻细胞丙二醛含量,说明青蒿素还能破坏细胞膜结构,造成胞内溶液外渗,致藻细胞死亡；但低浓度青蒿素适度提高绿藻超氧化物歧化酶和过氧化氢酶,有益于加速消除活性氧和自由基,减轻对绿藻细胞的危害。
在重庆市黄花蒿种植区,采用黄花蒿—茎瘤芥轮作种植模式,青蒿素的含量和产量、以及单位土地的年收入等显著高于黄花蒿—小麦和黄花蒿—油菜轮作种植模式。施用有机肥,尤其是有机无机适量配施显著促进黄花蒿生长,提高青蒿素含量和产量,减轻种植黄花蒿对土壤细菌、真菌和放线菌数量的抑制作用,提高土壤脲酶、蔗糖酶和磷酸酶活性,促进黄花蒿养分吸收。在干旱条件下,黄花蒿生长受到抑制,生物量的降幅高达69.36%,青蒿素产量可降低81.56%;同时黄花蒿叶片细胞膜透性增加,叶片水势、叶绿素含量,根系活力和氮、磷、钾吸收量显著降低。但轻度干旱提高了叶片青蒿素和根系磷、钾含量,故施用磷钾肥有益于提高黄花蒿抗旱性。此外,干旱使黄花蒿叶片脯氨酸含量增加了10余倍,增幅远远高于叶绿素、根系活力、养分吸收量和青蒿素含量的减少,说明脯氨酸积累可视为黄花蒿对干早产生的一种保护性生化反应。在黄花蒿种植过程中,移栽前炼苗促根和提高脯氨酸有益提高黄花蒿抗旱性。
综上所述,在重庆市黄花蒿种植区,友好栽培的内容包括采用黄花蒿—茎瘤芥轮作种植模式,适度延迟播种和移栽后茬作物,提倡施用有机肥,加强水分管理,分散种植,精细采收等。
Artemisia annua L., also called sweet wormwood, is an annual or biennial herbaceous plant used as a sole material for artemisinin extraction. Around40%of the populations in the world are now facing malaria infection. Artemisinin is recommended by World Health Organization (WHO) as a drug for the initial treatment of malaria.
About90%of A. annua and artemisinin in the world was grown and produced in China. Chongqing is the place for growing A. annua with high-quality and the yield of artemisinin and planting area of A. annua accounted for70%and80%, respectively, in China. Artemisinin is one of the allelochemicals effused from A. annua which could exist in soils for a long time, which could reduce land productivity, inhibit the growth of surrounding plants, and damage soil and water ecsystems.In the process of artificial cutivation of A. annua, artemisinin released into environments increased significantly with contents and yield of artemisinin increased, resulting in more serious allelopathic effect. Through field, quadrate survey, solution culture, greeenhouse pot and field experiments, we investigated colony characters of wild A. annua, species and growth of surrounding plants, the allelopathic effect and mechanism of artemisinin on terrestrial and aquatic plants. The influence of cropping pattern, fertilization and water management on A. annua itself and after-crops were also studied. Plant growth, concentration and yield of artemisinin were recorded in field and greenhouse pot experiments. The cultivation techniques for high yield of artemisinin and efficient soil utilization were primarily developed on the base of our studies. The main results are as follows:
The investigation of wild A. annua colonies in9southern provinces and cities with48samples showed that A. annua colonies were relatively stable.In the highest frequency of the9species pairs among different communities of wild A. annua,5species were Compositae,3Gramineae, and they were low connective. Few species of Cruciferae and Cyphomandra betacea were observed in or around the communities of wild A. annua, indicating allelopathic effects of A. annua on those plants, which inhibited significantly their growth and development.
Adding artemisinin produced by the A. annua into soils inhibited growth of after-crops such as wheat, rape and pickle. The biomass decreased by41.1～51.7%. The growth inhibition of pickle was much lower than rape and wheat. The inhibition of artemisinin on the growth of other plants was beneficial to enlarge living space of A. annua, by competition. Selection of after-crops with strong anti-allelopathic effect could increase whole soil productivity by improving land utilization. The high artemisinin reduced chlorophyll but increased carotenoid in the leaves of after-crops, indicating that artemisinin might promote chlorophyll decomposition, reduce photosynthetic rate, and decrease dry matter accumulation.In addition, artemisinin significantly inhibited the absorption of nitrogen, phosphorus and potassium, which might be related to the decreased root and nitrate reductase activities. Moderately late sowing and transplanting of after-crops following A. annua harvest might reduce the harm to the succeeding crop.
In solution culture, artemisinin inhibit the growth of green algae, and the density was reduced by74.5%(Chlorella pyrenoidosa) and61.8%(Scenedesmus obliquus), respectively. The decease in dissolved oxygen, chlorophyll and protein in green algae indicated that the poor growth might be due to slow photosynthesis and disorder protein synthesis caused by artemisinin. Cultivation of A. annua in large scale might reduce self-purification ability and productivity of water systems, leading to low pollution load and low fish production even their disappearances. Dispersive cultivation and fine recovery of A. annua at harvest could reduce artemisinin in water in the growing areas and eliminate the ecological risk.In addition, artemisinin increased the electric conductivity of culture medium and malondialdehyde in green algae cells, showing cell membrane damage, efflux of intracellular solution, and cell death by adding artemisinin into culture solutions. Low concentrations of artemisinin, however, increased the activities of superoxide dismutase, peroxide, and catalase in green algae, beneficial to fast elimination of active oxygen and free radicals, which could thus reduce the hazards to green algae cells.
In A. annua planting areas in Chongqing, the content and yield of artemisinin and the annual income per land unit were significantly higher under the cropping pattern of A. annua-B.juncea than A. annua-T aestivum L. and A. annua-B. napus L. Organic fertilization, especially organic and inorganic fertilizers, significantly promoted the growth of A. annua, improved the artemisinin content and yield, reduced the inhibitory effect of A annua on bacteria, fungi and actinomycetes in soils. Meanwhile, the activities of urease, invertase and phosphatasein soils with A. annua grown were increased and nutrient absorption by A. annua was also improved. Under drought stress, the biomass of A. annua was reduced by69.36%and artemisinin production by81.56%.Simultaneously, cell membrane permeability of leaves increased in contrast to leaf water potential, chlorophyll, root activity, and nutrient absorption (N, P, K) which were reduced significantly. Under mild drought stress, however, increased content of artemisinin in leaves and phosphorus and potassium in roots suggested that more phosphorus and potassium fertilizers might be beneficial to improve the drought resistance of A. annua. In addition, the proline in A. annua increased10times at least under drought stress, much more than the increament of chlorophyll, root activity, nutrient uptake, and reduction of artemisinin content. It seems reasonable to indicate that proline accumulation might be a protective biochemical reaction of A. annua under drought stress. In A. annua cultivation process, moderate drought stress applied to seedlings before transplanting might increase proline, beneficial to A. annua. resistant to drought.
In summary, in the A. annua planting area in Chongqing, friendly cultivation includes the cropping patterns of A. annua-B.juncea, moderate late sowing and transplanting of after-crops, organic fertilization, water management, disperse cultivation, and fine recovery at harvest.
我国黄花蒿种植面积和青蒿素产量约占全世界的90%以上,重庆市是中国黄花蒿的优质产区,种植面积和青蒿素产量分别占全国的70%和80%。青蒿素是黄花蒿(?)最多,在土壤中存在时间最长的化感物质,降低土地生产力,抑制周围和后茬植物生长,危害土壤和水体生态环境。在人工种植黄花蒿的过程中,随着青蒿素的含量和产量的提高,释放到环境中青蒿素大幅度增加,化感危害更加严重。论文采用生态学野外样方调查、培养试验、盆栽试验和田间试验等方法,研究了野生黄花蒿的群落特征及周围植物的存在状况,青蒿素对陆生和水生植物的化感效应及机理,轮作种植模式对后茬作物及自身的影响、施肥技术和水分管理对黄花蒿生长、青蒿素含量和产量,初步形成了青蒿素高产、土地利用高效的友好栽培技术。主要结果如下：
对南方9个省市48个样地野生黄花蒿种群调查发现,黄花蒿群落处于相对稳定状态。在野生黄花蒿不同群落中,出现频率最高的9个种对中,5种为菊科植物,3种禾本科植物,联结性较低。在野生黄花蒿群落中和周围,十字花科和茄科等植物极少,表明黄花蒿对其他植物具有化感效应,抑制植物的生长发育。
在土壤中添加黄花蒿产生的化感物质—青蒿素,不同程度抑制小麦、油菜和茎瘤芥等3种后茬作物的生长,生物量降幅高达41.1-51.7%,且对茎瘤芥的抑制作用低于油菜和小麦。青蒿素抑制其它植物生长,有利于扩大黄花蒿生存空间,增强竞争优势。但是,在黄花蒿种植区,选择抗化感作用较强的后茬作物可提高土地利用率和整体生产力。高浓度的青蒿素降低后茬作物叶绿素但提高类胡萝卜素含量,说明青蒿素促进叶绿素分解,导致光合速率下降,干物质积累减少。此外,青蒿素显著抑制氮、磷、钾养分吸收,与根系活力和硝酸还原酶活性降低有关。在黄花蒿收获后,适度延迟播种和移栽后茬作物可以减轻对后茬作物的危害。
在人工培养液中,不同浓度青蒿素显著抑制绿藻生长,绿藻密度可降低74.5%(蛋白核小球藻)和61.8%(斜生栅藻)。青蒿素使绿藻培养液溶解氧减少,绿藻叶绿素和蛋白质含量降低,说明青蒿素抑制绿藻生长与光合作用降低,蛋白合成受到干扰有关,大规模种植黄花蒿会降低水体自净能力和生产力,加剧污染负荷,鱼类减产和灭绝等；但分散种植和精细采收,有益于降低水体青蒿素浓度,减轻水环境风险。此外,青蒿素提高培养液电导率和绿藻细胞丙二醛含量,说明青蒿素还能破坏细胞膜结构,造成胞内溶液外渗,致藻细胞死亡；但低浓度青蒿素适度提高绿藻超氧化物歧化酶和过氧化氢酶,有益于加速消除活性氧和自由基,减轻对绿藻细胞的危害。
在重庆市黄花蒿种植区,采用黄花蒿—茎瘤芥轮作种植模式,青蒿素的含量和产量、以及单位土地的年收入等显著高于黄花蒿—小麦和黄花蒿—油菜轮作种植模式。施用有机肥,尤其是有机无机适量配施显著促进黄花蒿生长,提高青蒿素含量和产量,减轻种植黄花蒿对土壤细菌、真菌和放线菌数量的抑制作用,提高土壤脲酶、蔗糖酶和磷酸酶活性,促进黄花蒿养分吸收。在干旱条件下,黄花蒿生长受到抑制,生物量的降幅高达69.36%,青蒿素产量可降低81.56%;同时黄花蒿叶片细胞膜透性增加,叶片水势、叶绿素含量,根系活力和氮、磷、钾吸收量显著降低。但轻度干旱提高了叶片青蒿素和根系磷、钾含量,故施用磷钾肥有益于提高黄花蒿抗旱性。此外,干旱使黄花蒿叶片脯氨酸含量增加了10余倍,增幅远远高于叶绿素、根系活力、养分吸收量和青蒿素含量的减少,说明脯氨酸积累可视为黄花蒿对干早产生的一种保护性生化反应。在黄花蒿种植过程中,移栽前炼苗促根和提高脯氨酸有益提高黄花蒿抗旱性。
综上所述,在重庆市黄花蒿种植区,友好栽培的内容包括采用黄花蒿—茎瘤芥轮作种植模式,适度延迟播种和移栽后茬作物,提倡施用有机肥,加强水分管理,分散种植,精细采收等。
Artemisia annua L., also called sweet wormwood, is an annual or biennial herbaceous plant used as a sole material for artemisinin extraction. Around40%of the populations in the world are now facing malaria infection. Artemisinin is recommended by World Health Organization (WHO) as a drug for the initial treatment of malaria.
About90%of A. annua and artemisinin in the world was grown and produced in China. Chongqing is the place for growing A. annua with high-quality and the yield of artemisinin and planting area of A. annua accounted for70%and80%, respectively, in China. Artemisinin is one of the allelochemicals effused from A. annua which could exist in soils for a long time, which could reduce land productivity, inhibit the growth of surrounding plants, and damage soil and water ecsystems.In the process of artificial cutivation of A. annua, artemisinin released into environments increased significantly with contents and yield of artemisinin increased, resulting in more serious allelopathic effect. Through field, quadrate survey, solution culture, greeenhouse pot and field experiments, we investigated colony characters of wild A. annua, species and growth of surrounding plants, the allelopathic effect and mechanism of artemisinin on terrestrial and aquatic plants. The influence of cropping pattern, fertilization and water management on A. annua itself and after-crops were also studied. Plant growth, concentration and yield of artemisinin were recorded in field and greenhouse pot experiments. The cultivation techniques for high yield of artemisinin and efficient soil utilization were primarily developed on the base of our studies. The main results are as follows:
The investigation of wild A. annua colonies in9southern provinces and cities with48samples showed that A. annua colonies were relatively stable.In the highest frequency of the9species pairs among different communities of wild A. annua,5species were Compositae,3Gramineae, and they were low connective. Few species of Cruciferae and Cyphomandra betacea were observed in or around the communities of wild A. annua, indicating allelopathic effects of A. annua on those plants, which inhibited significantly their growth and development.
Adding artemisinin produced by the A. annua into soils inhibited growth of after-crops such as wheat, rape and pickle. The biomass decreased by41.1～51.7%. The growth inhibition of pickle was much lower than rape and wheat. The inhibition of artemisinin on the growth of other plants was beneficial to enlarge living space of A. annua, by competition. Selection of after-crops with strong anti-allelopathic effect could increase whole soil productivity by improving land utilization. The high artemisinin reduced chlorophyll but increased carotenoid in the leaves of after-crops, indicating that artemisinin might promote chlorophyll decomposition, reduce photosynthetic rate, and decrease dry matter accumulation.In addition, artemisinin significantly inhibited the absorption of nitrogen, phosphorus and potassium, which might be related to the decreased root and nitrate reductase activities. Moderately late sowing and transplanting of after-crops following A. annua harvest might reduce the harm to the succeeding crop.
In solution culture, artemisinin inhibit the growth of green algae, and the density was reduced by74.5%(Chlorella pyrenoidosa) and61.8%(Scenedesmus obliquus), respectively. The decease in dissolved oxygen, chlorophyll and protein in green algae indicated that the poor growth might be due to slow photosynthesis and disorder protein synthesis caused by artemisinin. Cultivation of A. annua in large scale might reduce self-purification ability and productivity of water systems, leading to low pollution load and low fish production even their disappearances. Dispersive cultivation and fine recovery of A. annua at harvest could reduce artemisinin in water in the growing areas and eliminate the ecological risk.In addition, artemisinin increased the electric conductivity of culture medium and malondialdehyde in green algae cells, showing cell membrane damage, efflux of intracellular solution, and cell death by adding artemisinin into culture solutions. Low concentrations of artemisinin, however, increased the activities of superoxide dismutase, peroxide, and catalase in green algae, beneficial to fast elimination of active oxygen and free radicals, which could thus reduce the hazards to green algae cells.
In A. annua planting areas in Chongqing, the content and yield of artemisinin and the annual income per land unit were significantly higher under the cropping pattern of A. annua-B.juncea than A. annua-T aestivum L. and A. annua-B. napus L. Organic fertilization, especially organic and inorganic fertilizers, significantly promoted the growth of A. annua, improved the artemisinin content and yield, reduced the inhibitory effect of A annua on bacteria, fungi and actinomycetes in soils. Meanwhile, the activities of urease, invertase and phosphatasein soils with A. annua grown were increased and nutrient absorption by A. annua was also improved. Under drought stress, the biomass of A. annua was reduced by69.36%and artemisinin production by81.56%.Simultaneously, cell membrane permeability of leaves increased in contrast to leaf water potential, chlorophyll, root activity, and nutrient absorption (N, P, K) which were reduced significantly. Under mild drought stress, however, increased content of artemisinin in leaves and phosphorus and potassium in roots suggested that more phosphorus and potassium fertilizers might be beneficial to improve the drought resistance of A. annua. In addition, the proline in A. annua increased10times at least under drought stress, much more than the increament of chlorophyll, root activity, nutrient uptake, and reduction of artemisinin content. It seems reasonable to indicate that proline accumulation might be a protective biochemical reaction of A. annua under drought stress. In A. annua cultivation process, moderate drought stress applied to seedlings before transplanting might increase proline, beneficial to A. annua. resistant to drought.
In summary, in the A. annua planting area in Chongqing, friendly cultivation includes the cropping patterns of A. annua-B.juncea, moderate late sowing and transplanting of after-crops, organic fertilization, water management, disperse cultivation, and fine recovery at harvest.
引文
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