当归连作障碍机制及其生物修复措施研究
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摘要
当归是常用中药材,但由于有限的适生区与市场周年需求的矛盾,导致当归连作现象普遍,连作障碍严重。2006-2008年,在当归主产区岷县清水乡布置田间和模拟试验,以不同连作茬次当归为研究对象,在从根际微生态和自毒作用角度揭示当归连作障碍主要原因的基础上,采用盆栽试验研究土壤生物因素和化感物质对当归生长过程中叶片保护酶和土壤根际微生态的影响,探讨当归连作障碍作用机制。最后,利用作物的化感效应,探索间作减缓当归连作障碍的可行性,得到如下主要结果:
1.连作通过降低植株保护酶活性引起活性氧代谢失调,导致活性氧积累和膜脂过氧化损伤。这种影响的外在表现是植株光合色素含量降低,光合作用强度下降,干物质积累减少,可溶性糖含量升高,生长受抑,病害严重,产量和品质下降。当归根膨大期,连茬当归叶片保护酶SOD、POD和CAT活性分别较正茬降低29.13%、10.87%和43.83%;渗透调节物质脯氨酸和可溶性糖含量分别增加121.95%和29.20%;膜脂过氧化的最终产物MDA含量增加16.00%;叶绿素a、叶绿素b和光合速率分别降低39.78%、46.90%和34.23%。株高、主根长、地上部和地下部干物重分别降低20.50%、61.50%、25.85%和35.94%,均达到显著差异。连作当归药材产量、挥发油含量及藁本内酯含量分别较正茬降低29.52%、28.42%和58.82%,而麻口病发病率是正茬的2.52倍。
2.当归连作导致土壤中可培养微生物数量减少,其中细菌和放线菌数量减少,而真菌数量明显增加,土壤微生物多样性指数减小。就整个生育时期平均而言,细菌、放线菌数量和多样性指数分别较正茬降低37.22%、43.68%和32.09%,而真菌数量增加106.64%。连作导致好气性纤维素分解菌和无机溶磷菌数量明显下降,氨化细菌数量大幅增加,而有机溶磷菌和钾细菌数量变化随生长时期不同而异。连作导致功能类群群落多样性和均匀度明显降低,优势集中性表现明显。土壤酶活性测定表明,连作土壤中脲酶和中性磷酸酶活性明显低于正茬,而过氧化氢酶和多酚氧化酶活性略有上升。
3.当归植株及根际土壤浸提液均具有生长抑制作用。浸提液的作用效果一般表现为低浓度不具有抑制作用,甚至具有促进作用,但随浓度升高抑制作用也随之增加。其中当归地上部浸提液较地下部浸提液的抑制作用更强,且根膨大期各种浸提液较苗期浸提液的抑制作用强。
4.连作土灭菌能在较大程度上改善当归生长,但不能完全消除连作带来的负效应。灭菌土上生长的当归株高、产量、挥发油含量和浸出物含量均高于连作土。当归叶片SOD和POD活性在灭菌土显著高于连作土,而CAT和MDA显著低于连作土。至收获期,灭菌土的细菌数量较连作土增加173.82%,而放线菌和真菌数量降低63.23%和37.87%;连作土灭菌处理改变土壤各功能类群的数量、多度、群落多样性、均匀度和优势集中性等特征参数,灭菌土的好气性纤维素分解菌、无机溶磷菌、钾细菌多度发生了变化(趋向于正茬),根膨大期和收获期其群落多样性分别较连作土提高65.51%和27.51%。灭菌土上当归麻口病发病率较连作土降低75.01%。
5.当归连作障碍的发生及加重是化感物质(T_1)和土壤生物因素(T_2)共同作用的结果。连作土壤中的化感物质和有害生物通过改变幼苗生理生化特性及其生长的微环境而使植株生长受阻,植株黄化,生物量下降,病虫害严重等,进而导致产量和品质下降。当归生长初期土壤中化学物质对其生长的作用强于土壤生物因素,而生长后期生物因素的作用强于化学物质,表明连作土壤中的化学物质不仅直接影响当归的生长,可能还通过影响土壤微生物而产生间接效应。
与对照相比,苗期当归地上部干重的减少中,连作土壤中化感物质和生物因素所占比例依次为61.90%和38.10%,地下部干重的减少中,二者所占比例依次为80.00%和20.00%;根膨大期地上部干重的减少中,二者所占比例依次为58.97%和41.03%,地下部干重的减少中,二者所占比例依次为41.02%和58.98%;当归收获期,药材产量的减少中,二者所占比例依次为48.29%和51.71%;挥发油含量下降值中,二者所占比例依次为21.74%和78.26%;浸出物含量下降值中,二者所占比例依次为47.16%和52.84%。
连作土壤中化感物质的加入使当归叶片SOD、POD和CAT活性分别较CK增加12.50%、60.47%和6.43%,其中POD达到显著差异,MDA含量增加14.67%,差异显著;在化感作用的基础上,连作土壤中生物因素的加入使当归叶片SOD和CAT活性分别较CK降低20.59%和28.22%,POD活性和MDA含量分别较CK增加38.22%和32.00%,差异均显著。
与对照相比,T_1和T_2处理均对真菌生长繁殖有显著促进作用,对放线菌、氨化细菌、好气性纤维素分解菌、有机溶磷菌和钾细菌有明显抑制作用。T_1和T_2处理根膨大期均可显著降低细菌数量,增加无机溶磷菌数量,而收获期则显著增加细菌数量,降低无机溶磷菌数量。根膨大期T_1和T_2处理当归根际土壤好气性纤维素分解菌多度发生了变化,收获期T_1处理好气性纤维素分解菌多度发生了变化,T_2处理钾细菌多度发生变化。T_1和T_2处理功能类群总数、群落多样性和均匀度均明显下降,优势集中性表现明显,且根膨大期多样性参数值以及功能类群数量的变化幅度大于收获期。与CK相比,根膨大期T_1处理功能类群总数和群落多样性分别降低1.92%和21.97%,T_2处理分别降低25.13%和43.88%。
6.大蒜与当归间作显著改善连作土壤中微生物区系,提高功能类群多样性及均匀度,丰富了土壤微生物群落结构,抑制和影响病原菌的生长、减少病害的发生,植株体内保护酶活性提高,膜脂过氧化程度降低,从生理上提高当归自身的抗性,改善了当归连作障碍。
根膨大期,间作处理当归株高、地上干物重和地下干物重分别较单作提高12.34%、36.92%和9.15%。间作增强当归自身防御酶体系,与单作相比,间作SOD、POD和CAT活性分别提高80.00%、243.19%和37.13%,MDA含量降低11.67%。间作显著降低麻口病发病率,而提高当归药材产量和品质,与单作相比,间作当归药材麻口病发病率下降56.25%,产量、挥发油含量和浸出物含量分别增加52.77%、50.00%和10.20%。
间作有利于提高生长盛期土壤细菌、好气性纤维素分解菌、有机溶磷菌、无机溶磷菌和钾细菌数量。当归根膨大期,间作土壤细菌、好气性纤维素分解菌、有机溶磷菌、无机溶磷菌数量分别较单作提高53.52%、23.19%、103.44%和49.46%。收获期,间作显著降低细菌、真菌和氨化细菌数量,提高无机溶磷菌和钾细菌数量,各指标的增减幅度分别为-31.71%、-22.05%、-47.26%、20.05%和75.59%。间作条件下生长盛期当归根际土壤好气性纤维素分解菌、有机溶磷菌和钾细菌的多度发生变化。尽管功能类群总数间作低于单作处理,但群落多样性和均匀度均明显提高,优势集中性表现不明显,且根膨大期多样性参数值及功能类群数量的变化幅度大于苗期和收获期。间作功能类群多样性指数在苗期、根膨大期和收获期分别较单作提高11.25%、80.81%和44.97%
7.综合以上研究结果,认为当归连作障碍是由化感物质、土壤有害生物等多种因素综合作用的结果。土壤中化感物质积累,一方面可能导致自毒作用,引起产量和品质的下降;另一方面可能通过破坏根系的细胞膜,使病虫害易于侵入,或是通过改变土壤微生物区系,刺激某些病原菌的增殖,使病害发生严重,进而导致产量和品质下降。当归与大蒜间作可以通过改善土壤微生物区系、提高植株自身抗性,抑制病原菌生长,缓解当归连作障碍。
Angelica sisnensis is a perennial herb of Umbelliferae,and has been cultivated for more than 1700 years.Its root is used frequently as a crude drug in traditional Chinese medicine.However,the contradiction between limited suitable areas and market supply all year long resulted in the fact that continuous cropping obstacles become common and serious more and more,but its reason is not clear up to now.To explore the origin of its occurrence mechanisms,field and pot experiment were conducted in Min county of Gansu province during 2006-2008.
Regarding allelopathic substances and microorganisms in rhizosphere soil as the ligament of many factors of continuous cropping obstacles,we first detected the variation of rhizosphere microbial population in response to continuous A.sinensis cropping,then collected and identified the autotoxic effect of plant and soil arounding A.sinensis.In addition,we also trialed the effect of allelochemicals in continuous cropping soil on seedlings growth,rhizosphere microbial and Ditylenchus destructor,and the interactions of allelopathy and microbial from continuous cropping soil on A.sinensis growth.At last we trialed the effect of intercropping on A.sinensis growth and the microflora in rhizosphere soil of A.sinensis meanwhile try to supply the scientific bases for the alleviation or elimination of A.sinensis continuous cropping obstacles.The main results are as follows:
1.Through decreasing the activity of protective enzymes and their ability of cleaning free radicle, continuous cropping made lots of free radicle remain in plant so that induced membrane lipid peroxidization,electrolytic leakage became heavier,content of proline and soluble sugar increased.The external manifestation of this influences were that plant growth was inhibited,content of photosynthetic pigments decreased,so did the intensity of photosynthesis and respiration,content of dry mass.In continuous A.sinensis cropping system,SOD,POD and CAT activity decreased by 29.13%,10.87%and 43.83%respectively compared with those in non-replant A.sinensis;The content of proline,soluble sugar and MDA increased by 121.95%,29.20%and 16.00%;The content of chlorophyll a,chlorophyll b and photosynthetic rate decreased by 39.78%,46.90%and 34.23%respectively;The yield,essential oil content and extract content decreased by 29.52%,28.42%and 58.82%respectively,and the incidence of D.destructor increased by 2.52 fold.Both of the decrease or increase extent showed significance at P＜0.05.
2.Plate count analysis revealed a reduction trend in the number of culturable microorganism in rhizosphere as A.sinensis cropping continued.Bacteria and actinomyces had an apparent decrease extent in CFU number,while fungi had an apparent increase extent.In the second cropping season,the number of culturable bacteria,actinomyces,and diversity index in rhizosphere soil were decreased by 37.22%, 43.68%and 32.09%respectively compared with those in non-replant A.sinensis soil,while the number of fungi was increased by 106.64%.An obvious decrease tread was also found in the number of aerobic cellulose-decomposing bacteria and inorganic phosphorus-solubilizing bacteria,while great increase in ammonifying bacteria.The variation of organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria changed with the change of growth stage.Of the essential soil enzymes,urease and neutral phosphatase activity significantly declined in the continuous cropping,but the catalase and polyphenol oxidase activity rise little.
3.All of the aqueous extracts from A.sinensis tissues and rhizosphere soil had inhibitory effects on A.sinensis germination and seedling growth,and this inhibitory effect generally increased with increased extract concentration at a certain ranges of extract concentrations.Under lower concentration,the inhibition wasn't shown,indeed,the promotion was exhibited.With the concentration increasing,the inhibition was enhanced.Inhibitory effects of plant above ground extracts were more evident than that of root extracts.Inhibitory effects were greater for all extracts at the rootstock thickening than the seedling stage.It may be concluded that soil previously planted with A.sinensis as well as A.sinensis plants themselves contain allelopathic substances.They are toxic to the growth of A.sinensis and other plants. The chemical compounds may play some part in growth inhibition due to the continuous cropping of A. sinensis.
4.Soil sterilization could improve growth of A.sinensis plants to greater extent and for example, plant height,yield,essential oil content and extract content of A.sinensis grown in sterilized soil were significantly higher than those in non-sterilized soil.SOD and POD activities in the leaves of A.sinensis plants grown in sterilized soil were higher than those in non-sterilized soil,while CAT activity and MDA content were significantly lower.The number of bacteria in sterilized soil was increased by 173.82% compared with that in non-sterilized soil,while the number of actinomyces and fungi were decreased by 63.23%and 37.87%respectively.The number,abundance,community diversity,evenness and dominance concentration of some functional groups were changed greatly in sterilized soil.The abundance of aerobic cellulose-decomposing bacteria,inorganic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in sterilized soil had changed to tends to those in rotation system.The diversity index in sterilized soil decreased by 31.04%at seedling stage,but increased by 65.51%and 27.51%at rootstock thickening and harvest stage compared with those in non-sterilized soil.The change range of diversity index and population of functional groups in rhizosphere soils at the rootstock thickening was higher than at seedling and harvest stages.The incidence of D.destructor of A.sinensis grown in sterilized soil decreased by 75.01%compared with that in non-sterilized soil.Although there are more improving in factors affected growth of A.sinensis,the A.sinensis continuous cropping obstacle had not been eliminated completely by soil sterilization.
5.The occurrence and aggravation of A.sinensis continuous cropping obstacle was resulted from both allelopathy and soil biota factors.The growth,yield and quality of A.sinensis plants were affected by the allelopathy and the soil biota which in continuous cropping soil through the change of plant physiological and biochemical characteristics and soil microenvironment of plant growth.The dry weight of A.sinensis was significantly depressed in the treatments other than in the control,and the inhibitory effect of allelopathy on growth of A.sinensis was higher than that of deleterious microorganisms in early growth period,whereas the inhibitory effect showed a contrary tendency in late growth period.
Compared to the control,for the inhibition percentage of biomass,the proportion of T_1 treatment and T_2 treatment was 61.90%and 38.10%for shoot dry weight,80.00%and 20.00%for root dry weight at seedling stage;The proportion of T_1 treatment and T_2 treatment was 58.97%and 41.03%for shoot dry weight,41.02%and 58.98%for root dry weight at rootstock thickening.The percentage was 48.29%and 51.71%for yield,21.74%and 78.26%for essential oil content,47.16%and 2.84%for extract content respectively.
For the protective enzymatic activity,compared to the control,SOD,POD and CAT activity were increased by 12.50%,60.47%and 6.43%in the leaves of A.sinensis plants as the T_1 treatment,MDA content was increased by 14.67%.SOD and CAT activity were decreased by 20.59%and 28.22%,while POD activity and MDA content were increased by 38.22%and 32.00%as the T_2 treatment,and the differences were significant all.
The number,abundance,community diversity,evenness and dominance concentration of some functional proups also were changed markedly in the effect of allelopathic compounds and deleterious microorganisms in replant soil.Compared with the control,allelopathic compounds and deleterious microorganisms in the replant soil seemed to stimulate the reproduction of rhizosphere soil fungi,and restrain the reproduction of rhizosphere soil actinomyces,ammonifying bacteria,aerobic cellulosedecomposing bacteria,organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria at rootstock thickening and harvest stage.T_1 and T_2 decreased the population of bacteria and increased that of inorganic phosphorus-solubilizing bacteria significantly at rootstock thickening,whereas they increased the population of bacteria and decreased that of inorganic phosphorus-solubilizing bacteria significantly at the harvest stage.The abundance of aerobic cellulose-decomposing bacteria and kalium-solubilizing bacteria in T_1 and T_2 A.sinensis rhizosphere soils changed at the rootstock thickening and harvest stage. The total population of functional groups,community diversity and evenness decreased and dominance concentration increased in T_1 and T_2 A.sinensis rhizosphere soils compared with the control.The change range of diversity index and population of functional groups in rhizosphere soils at the rootstock thickening was higher than at the harvest stage,compared to the control,The total population and community diversity of functional groups were decreased by 1.92%and 21.97%in T_1 and 25.13%, 43.88%in T_2 at the rootstock thickening.
6.The microflora in rhizosphere soil of A.sinensis was significantly improved in intercropping system by restraining pathogen growth and enhancing community diversity and evenness of some functional proups.Which alleviated the A.sinensis continuous cropping problem by restraining diseases and regulating the protective enzymes system.
A.sinensis plant growth status was improved in intercropping system.Compared with the monocropping system,plant height,shoot dry weight and root dry weight were increased by 12.34%, 36.92%and 9.15%respectively.SOD,POD and CAT activity were increased by 80.00%,243.19%and 37.13%respectively,while MDA content was decreased by 11.67%.Lower D.destructor incidence and higher yield and quality of A.sinensis were observed in intercropping system compared to those in monocropping system.
The population of bacteria,aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria,inorganic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in rhizosphere soil could be promoted in intercropping in vigorous growth stage.The number of bacteria,aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria and inorganic phosphorussolubilizing bacteria in intercropping were enhanced by 53.52%,23.19%,103.44%and 49.46% respectively compared with the monocropping.In harvest stage,the number of bacteria,fungi and ammonifying bacteria significantly decreased by 31.71%,22.05%and 47.26%,while the population of inorganic phosphorus-solubilizing bacteria,kalium-solubilizing bacteria increased by 20.05%and 75.59%. The abundance of aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in intercropping A.sinensis rhizosphere soils changed at the rootstock thickening.Although the total population of functional groups in intercropping A.sinensis rhizosphere soils was lower than in monocroping system,community diversity and evenness increased and dominance concentration decreased.The change range of diversity index and population of functional groups in intercropping A.sinensis rhizosphere soils at the rootstock thickening was higher than at the seedling and harvest stage.The diversity index was significantly increased by 11.25%,80.81%and 44.97% respectively at the seedling,rootstock thickening and harvest stage in intercropping system compared with the control.
7.Based on the experimental results described above,we concluded the allelopathic chemicals and deleterious microorganisms affected A.sinensis growth and cause continuous A.sinensis cropping problem. Why was the A.sinensis growth inhibited severely when it was continuously cropped in the same field? It is supposed that after the whole growth stage in the first year,the rhizosphere ecology was changed so greatly that it was no longer suitable for A.sinensis growth.One result was that a lot of allelochemicals were produced directly or indirectly in four ways including leaching from the shoot,shoot and root residues,root exudates and metabolites of microorganisms in rhizosphere,which will take effect immediately after seedling was cultivated in the following year.The other result indicated that the quality and quantity of microorganisms in the rhizosphere were changed greatly,including the deleterious microorganism of A.sinensis.In the second year,after cultivating,the allelopathic substances would be absorbed by A.sinensis.Some of the deleterious microorganisms would also take action at this time. Therefore,in the early stage the A.sinensis development would be retarded,caused low protective enzyme activity and shoot biomass.The plant developed in the condition will have weaker resistance and is easier for pathogens to infect than normal A.sinensis in shis stage.At the following stage,more pathogens will also infect A.sinensis and develop.During the process,they will cause the decline of yield and quality through change the physiological character.In a word,all the factors collectively caused the continuous A.sinensis cropping problem.In intercropping system,continuous A.sinensis cropping obstacle was significantly alleviated by changing the microflora in rhizosphere soil and regulating the protective enzymes system of A.sinensis.
1.连作通过降低植株保护酶活性引起活性氧代谢失调,导致活性氧积累和膜脂过氧化损伤。这种影响的外在表现是植株光合色素含量降低,光合作用强度下降,干物质积累减少,可溶性糖含量升高,生长受抑,病害严重,产量和品质下降。当归根膨大期,连茬当归叶片保护酶SOD、POD和CAT活性分别较正茬降低29.13%、10.87%和43.83%;渗透调节物质脯氨酸和可溶性糖含量分别增加121.95%和29.20%;膜脂过氧化的最终产物MDA含量增加16.00%;叶绿素a、叶绿素b和光合速率分别降低39.78%、46.90%和34.23%。株高、主根长、地上部和地下部干物重分别降低20.50%、61.50%、25.85%和35.94%,均达到显著差异。连作当归药材产量、挥发油含量及藁本内酯含量分别较正茬降低29.52%、28.42%和58.82%,而麻口病发病率是正茬的2.52倍。
2.当归连作导致土壤中可培养微生物数量减少,其中细菌和放线菌数量减少,而真菌数量明显增加,土壤微生物多样性指数减小。就整个生育时期平均而言,细菌、放线菌数量和多样性指数分别较正茬降低37.22%、43.68%和32.09%,而真菌数量增加106.64%。连作导致好气性纤维素分解菌和无机溶磷菌数量明显下降,氨化细菌数量大幅增加,而有机溶磷菌和钾细菌数量变化随生长时期不同而异。连作导致功能类群群落多样性和均匀度明显降低,优势集中性表现明显。土壤酶活性测定表明,连作土壤中脲酶和中性磷酸酶活性明显低于正茬,而过氧化氢酶和多酚氧化酶活性略有上升。
3.当归植株及根际土壤浸提液均具有生长抑制作用。浸提液的作用效果一般表现为低浓度不具有抑制作用,甚至具有促进作用,但随浓度升高抑制作用也随之增加。其中当归地上部浸提液较地下部浸提液的抑制作用更强,且根膨大期各种浸提液较苗期浸提液的抑制作用强。
4.连作土灭菌能在较大程度上改善当归生长,但不能完全消除连作带来的负效应。灭菌土上生长的当归株高、产量、挥发油含量和浸出物含量均高于连作土。当归叶片SOD和POD活性在灭菌土显著高于连作土,而CAT和MDA显著低于连作土。至收获期,灭菌土的细菌数量较连作土增加173.82%,而放线菌和真菌数量降低63.23%和37.87%;连作土灭菌处理改变土壤各功能类群的数量、多度、群落多样性、均匀度和优势集中性等特征参数,灭菌土的好气性纤维素分解菌、无机溶磷菌、钾细菌多度发生了变化(趋向于正茬),根膨大期和收获期其群落多样性分别较连作土提高65.51%和27.51%。灭菌土上当归麻口病发病率较连作土降低75.01%。
5.当归连作障碍的发生及加重是化感物质(T_1)和土壤生物因素(T_2)共同作用的结果。连作土壤中的化感物质和有害生物通过改变幼苗生理生化特性及其生长的微环境而使植株生长受阻,植株黄化,生物量下降,病虫害严重等,进而导致产量和品质下降。当归生长初期土壤中化学物质对其生长的作用强于土壤生物因素,而生长后期生物因素的作用强于化学物质,表明连作土壤中的化学物质不仅直接影响当归的生长,可能还通过影响土壤微生物而产生间接效应。
与对照相比,苗期当归地上部干重的减少中,连作土壤中化感物质和生物因素所占比例依次为61.90%和38.10%,地下部干重的减少中,二者所占比例依次为80.00%和20.00%;根膨大期地上部干重的减少中,二者所占比例依次为58.97%和41.03%,地下部干重的减少中,二者所占比例依次为41.02%和58.98%;当归收获期,药材产量的减少中,二者所占比例依次为48.29%和51.71%;挥发油含量下降值中,二者所占比例依次为21.74%和78.26%;浸出物含量下降值中,二者所占比例依次为47.16%和52.84%。
连作土壤中化感物质的加入使当归叶片SOD、POD和CAT活性分别较CK增加12.50%、60.47%和6.43%,其中POD达到显著差异,MDA含量增加14.67%,差异显著;在化感作用的基础上,连作土壤中生物因素的加入使当归叶片SOD和CAT活性分别较CK降低20.59%和28.22%,POD活性和MDA含量分别较CK增加38.22%和32.00%,差异均显著。
与对照相比,T_1和T_2处理均对真菌生长繁殖有显著促进作用,对放线菌、氨化细菌、好气性纤维素分解菌、有机溶磷菌和钾细菌有明显抑制作用。T_1和T_2处理根膨大期均可显著降低细菌数量,增加无机溶磷菌数量,而收获期则显著增加细菌数量,降低无机溶磷菌数量。根膨大期T_1和T_2处理当归根际土壤好气性纤维素分解菌多度发生了变化,收获期T_1处理好气性纤维素分解菌多度发生了变化,T_2处理钾细菌多度发生变化。T_1和T_2处理功能类群总数、群落多样性和均匀度均明显下降,优势集中性表现明显,且根膨大期多样性参数值以及功能类群数量的变化幅度大于收获期。与CK相比,根膨大期T_1处理功能类群总数和群落多样性分别降低1.92%和21.97%,T_2处理分别降低25.13%和43.88%。
6.大蒜与当归间作显著改善连作土壤中微生物区系,提高功能类群多样性及均匀度,丰富了土壤微生物群落结构,抑制和影响病原菌的生长、减少病害的发生,植株体内保护酶活性提高,膜脂过氧化程度降低,从生理上提高当归自身的抗性,改善了当归连作障碍。
根膨大期,间作处理当归株高、地上干物重和地下干物重分别较单作提高12.34%、36.92%和9.15%。间作增强当归自身防御酶体系,与单作相比,间作SOD、POD和CAT活性分别提高80.00%、243.19%和37.13%,MDA含量降低11.67%。间作显著降低麻口病发病率,而提高当归药材产量和品质,与单作相比,间作当归药材麻口病发病率下降56.25%,产量、挥发油含量和浸出物含量分别增加52.77%、50.00%和10.20%。
间作有利于提高生长盛期土壤细菌、好气性纤维素分解菌、有机溶磷菌、无机溶磷菌和钾细菌数量。当归根膨大期,间作土壤细菌、好气性纤维素分解菌、有机溶磷菌、无机溶磷菌数量分别较单作提高53.52%、23.19%、103.44%和49.46%。收获期,间作显著降低细菌、真菌和氨化细菌数量,提高无机溶磷菌和钾细菌数量,各指标的增减幅度分别为-31.71%、-22.05%、-47.26%、20.05%和75.59%。间作条件下生长盛期当归根际土壤好气性纤维素分解菌、有机溶磷菌和钾细菌的多度发生变化。尽管功能类群总数间作低于单作处理,但群落多样性和均匀度均明显提高,优势集中性表现不明显,且根膨大期多样性参数值及功能类群数量的变化幅度大于苗期和收获期。间作功能类群多样性指数在苗期、根膨大期和收获期分别较单作提高11.25%、80.81%和44.97%
7.综合以上研究结果,认为当归连作障碍是由化感物质、土壤有害生物等多种因素综合作用的结果。土壤中化感物质积累,一方面可能导致自毒作用,引起产量和品质的下降;另一方面可能通过破坏根系的细胞膜,使病虫害易于侵入,或是通过改变土壤微生物区系,刺激某些病原菌的增殖,使病害发生严重,进而导致产量和品质下降。当归与大蒜间作可以通过改善土壤微生物区系、提高植株自身抗性,抑制病原菌生长,缓解当归连作障碍。
Angelica sisnensis is a perennial herb of Umbelliferae,and has been cultivated for more than 1700 years.Its root is used frequently as a crude drug in traditional Chinese medicine.However,the contradiction between limited suitable areas and market supply all year long resulted in the fact that continuous cropping obstacles become common and serious more and more,but its reason is not clear up to now.To explore the origin of its occurrence mechanisms,field and pot experiment were conducted in Min county of Gansu province during 2006-2008.
Regarding allelopathic substances and microorganisms in rhizosphere soil as the ligament of many factors of continuous cropping obstacles,we first detected the variation of rhizosphere microbial population in response to continuous A.sinensis cropping,then collected and identified the autotoxic effect of plant and soil arounding A.sinensis.In addition,we also trialed the effect of allelochemicals in continuous cropping soil on seedlings growth,rhizosphere microbial and Ditylenchus destructor,and the interactions of allelopathy and microbial from continuous cropping soil on A.sinensis growth.At last we trialed the effect of intercropping on A.sinensis growth and the microflora in rhizosphere soil of A.sinensis meanwhile try to supply the scientific bases for the alleviation or elimination of A.sinensis continuous cropping obstacles.The main results are as follows:
1.Through decreasing the activity of protective enzymes and their ability of cleaning free radicle, continuous cropping made lots of free radicle remain in plant so that induced membrane lipid peroxidization,electrolytic leakage became heavier,content of proline and soluble sugar increased.The external manifestation of this influences were that plant growth was inhibited,content of photosynthetic pigments decreased,so did the intensity of photosynthesis and respiration,content of dry mass.In continuous A.sinensis cropping system,SOD,POD and CAT activity decreased by 29.13%,10.87%and 43.83%respectively compared with those in non-replant A.sinensis;The content of proline,soluble sugar and MDA increased by 121.95%,29.20%and 16.00%;The content of chlorophyll a,chlorophyll b and photosynthetic rate decreased by 39.78%,46.90%and 34.23%respectively;The yield,essential oil content and extract content decreased by 29.52%,28.42%and 58.82%respectively,and the incidence of D.destructor increased by 2.52 fold.Both of the decrease or increase extent showed significance at P＜0.05.
2.Plate count analysis revealed a reduction trend in the number of culturable microorganism in rhizosphere as A.sinensis cropping continued.Bacteria and actinomyces had an apparent decrease extent in CFU number,while fungi had an apparent increase extent.In the second cropping season,the number of culturable bacteria,actinomyces,and diversity index in rhizosphere soil were decreased by 37.22%, 43.68%and 32.09%respectively compared with those in non-replant A.sinensis soil,while the number of fungi was increased by 106.64%.An obvious decrease tread was also found in the number of aerobic cellulose-decomposing bacteria and inorganic phosphorus-solubilizing bacteria,while great increase in ammonifying bacteria.The variation of organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria changed with the change of growth stage.Of the essential soil enzymes,urease and neutral phosphatase activity significantly declined in the continuous cropping,but the catalase and polyphenol oxidase activity rise little.
3.All of the aqueous extracts from A.sinensis tissues and rhizosphere soil had inhibitory effects on A.sinensis germination and seedling growth,and this inhibitory effect generally increased with increased extract concentration at a certain ranges of extract concentrations.Under lower concentration,the inhibition wasn't shown,indeed,the promotion was exhibited.With the concentration increasing,the inhibition was enhanced.Inhibitory effects of plant above ground extracts were more evident than that of root extracts.Inhibitory effects were greater for all extracts at the rootstock thickening than the seedling stage.It may be concluded that soil previously planted with A.sinensis as well as A.sinensis plants themselves contain allelopathic substances.They are toxic to the growth of A.sinensis and other plants. The chemical compounds may play some part in growth inhibition due to the continuous cropping of A. sinensis.
4.Soil sterilization could improve growth of A.sinensis plants to greater extent and for example, plant height,yield,essential oil content and extract content of A.sinensis grown in sterilized soil were significantly higher than those in non-sterilized soil.SOD and POD activities in the leaves of A.sinensis plants grown in sterilized soil were higher than those in non-sterilized soil,while CAT activity and MDA content were significantly lower.The number of bacteria in sterilized soil was increased by 173.82% compared with that in non-sterilized soil,while the number of actinomyces and fungi were decreased by 63.23%and 37.87%respectively.The number,abundance,community diversity,evenness and dominance concentration of some functional groups were changed greatly in sterilized soil.The abundance of aerobic cellulose-decomposing bacteria,inorganic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in sterilized soil had changed to tends to those in rotation system.The diversity index in sterilized soil decreased by 31.04%at seedling stage,but increased by 65.51%and 27.51%at rootstock thickening and harvest stage compared with those in non-sterilized soil.The change range of diversity index and population of functional groups in rhizosphere soils at the rootstock thickening was higher than at seedling and harvest stages.The incidence of D.destructor of A.sinensis grown in sterilized soil decreased by 75.01%compared with that in non-sterilized soil.Although there are more improving in factors affected growth of A.sinensis,the A.sinensis continuous cropping obstacle had not been eliminated completely by soil sterilization.
5.The occurrence and aggravation of A.sinensis continuous cropping obstacle was resulted from both allelopathy and soil biota factors.The growth,yield and quality of A.sinensis plants were affected by the allelopathy and the soil biota which in continuous cropping soil through the change of plant physiological and biochemical characteristics and soil microenvironment of plant growth.The dry weight of A.sinensis was significantly depressed in the treatments other than in the control,and the inhibitory effect of allelopathy on growth of A.sinensis was higher than that of deleterious microorganisms in early growth period,whereas the inhibitory effect showed a contrary tendency in late growth period.
Compared to the control,for the inhibition percentage of biomass,the proportion of T_1 treatment and T_2 treatment was 61.90%and 38.10%for shoot dry weight,80.00%and 20.00%for root dry weight at seedling stage;The proportion of T_1 treatment and T_2 treatment was 58.97%and 41.03%for shoot dry weight,41.02%and 58.98%for root dry weight at rootstock thickening.The percentage was 48.29%and 51.71%for yield,21.74%and 78.26%for essential oil content,47.16%and 2.84%for extract content respectively.
For the protective enzymatic activity,compared to the control,SOD,POD and CAT activity were increased by 12.50%,60.47%and 6.43%in the leaves of A.sinensis plants as the T_1 treatment,MDA content was increased by 14.67%.SOD and CAT activity were decreased by 20.59%and 28.22%,while POD activity and MDA content were increased by 38.22%and 32.00%as the T_2 treatment,and the differences were significant all.
The number,abundance,community diversity,evenness and dominance concentration of some functional proups also were changed markedly in the effect of allelopathic compounds and deleterious microorganisms in replant soil.Compared with the control,allelopathic compounds and deleterious microorganisms in the replant soil seemed to stimulate the reproduction of rhizosphere soil fungi,and restrain the reproduction of rhizosphere soil actinomyces,ammonifying bacteria,aerobic cellulosedecomposing bacteria,organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria at rootstock thickening and harvest stage.T_1 and T_2 decreased the population of bacteria and increased that of inorganic phosphorus-solubilizing bacteria significantly at rootstock thickening,whereas they increased the population of bacteria and decreased that of inorganic phosphorus-solubilizing bacteria significantly at the harvest stage.The abundance of aerobic cellulose-decomposing bacteria and kalium-solubilizing bacteria in T_1 and T_2 A.sinensis rhizosphere soils changed at the rootstock thickening and harvest stage. The total population of functional groups,community diversity and evenness decreased and dominance concentration increased in T_1 and T_2 A.sinensis rhizosphere soils compared with the control.The change range of diversity index and population of functional groups in rhizosphere soils at the rootstock thickening was higher than at the harvest stage,compared to the control,The total population and community diversity of functional groups were decreased by 1.92%and 21.97%in T_1 and 25.13%, 43.88%in T_2 at the rootstock thickening.
6.The microflora in rhizosphere soil of A.sinensis was significantly improved in intercropping system by restraining pathogen growth and enhancing community diversity and evenness of some functional proups.Which alleviated the A.sinensis continuous cropping problem by restraining diseases and regulating the protective enzymes system.
A.sinensis plant growth status was improved in intercropping system.Compared with the monocropping system,plant height,shoot dry weight and root dry weight were increased by 12.34%, 36.92%and 9.15%respectively.SOD,POD and CAT activity were increased by 80.00%,243.19%and 37.13%respectively,while MDA content was decreased by 11.67%.Lower D.destructor incidence and higher yield and quality of A.sinensis were observed in intercropping system compared to those in monocropping system.
The population of bacteria,aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria,inorganic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in rhizosphere soil could be promoted in intercropping in vigorous growth stage.The number of bacteria,aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria and inorganic phosphorussolubilizing bacteria in intercropping were enhanced by 53.52%,23.19%,103.44%and 49.46% respectively compared with the monocropping.In harvest stage,the number of bacteria,fungi and ammonifying bacteria significantly decreased by 31.71%,22.05%and 47.26%,while the population of inorganic phosphorus-solubilizing bacteria,kalium-solubilizing bacteria increased by 20.05%and 75.59%. The abundance of aerobic cellulose-decomposing bacteria,organic phosphorus-solubilizing bacteria and kalium-solubilizing bacteria in intercropping A.sinensis rhizosphere soils changed at the rootstock thickening.Although the total population of functional groups in intercropping A.sinensis rhizosphere soils was lower than in monocroping system,community diversity and evenness increased and dominance concentration decreased.The change range of diversity index and population of functional groups in intercropping A.sinensis rhizosphere soils at the rootstock thickening was higher than at the seedling and harvest stage.The diversity index was significantly increased by 11.25%,80.81%and 44.97% respectively at the seedling,rootstock thickening and harvest stage in intercropping system compared with the control.
7.Based on the experimental results described above,we concluded the allelopathic chemicals and deleterious microorganisms affected A.sinensis growth and cause continuous A.sinensis cropping problem. Why was the A.sinensis growth inhibited severely when it was continuously cropped in the same field? It is supposed that after the whole growth stage in the first year,the rhizosphere ecology was changed so greatly that it was no longer suitable for A.sinensis growth.One result was that a lot of allelochemicals were produced directly or indirectly in four ways including leaching from the shoot,shoot and root residues,root exudates and metabolites of microorganisms in rhizosphere,which will take effect immediately after seedling was cultivated in the following year.The other result indicated that the quality and quantity of microorganisms in the rhizosphere were changed greatly,including the deleterious microorganism of A.sinensis.In the second year,after cultivating,the allelopathic substances would be absorbed by A.sinensis.Some of the deleterious microorganisms would also take action at this time. Therefore,in the early stage the A.sinensis development would be retarded,caused low protective enzyme activity and shoot biomass.The plant developed in the condition will have weaker resistance and is easier for pathogens to infect than normal A.sinensis in shis stage.At the following stage,more pathogens will also infect A.sinensis and develop.During the process,they will cause the decline of yield and quality through change the physiological character.In a word,all the factors collectively caused the continuous A.sinensis cropping problem.In intercropping system,continuous A.sinensis cropping obstacle was significantly alleviated by changing the microflora in rhizosphere soil and regulating the protective enzymes system of A.sinensis.
引文
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