连作条件下马铃薯根际微生态环境的变化及其生物效应研究
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摘要
针对马铃薯生产中存在的因长期连作引起的病虫害严重以及产量持续下降等问题，本研究于2010年在甘肃景泰条山农场马铃薯种植区域内选择不同连作年限的相邻地块进行田间试验，以轮作地块为对照（即连作0年，用L0表示），其余依次为连作1年（L1）、2年（L2）、3年（L3）、4年（L4）和5年（L5），从根际微生态的角度出发，采用传统平板培养和PCR-DGGE指纹图谱技术相结合的方法，研究马铃薯连作条件下以根际微生物群落组成、根际细菌和真菌种群结构的变化、以及根系生理变化为特征的根际微生态环境的改变，以及根际微生态环境改变后对植株保护酶活性、丙二醛含量、植株主要生物学性状和产量的影响等生物效应，以揭示马铃薯连作障碍的可能机理，为马铃薯连作障碍的综合防控提供理论依据。主要研究结果如下：
1.连作对马铃薯根际微生态环境的影响
马铃薯根际可培养微生物的数量对连作的反应程度明显不同，细菌对连作的反应较敏感，放线菌反应稍滞后。随着连作年限的增加，马铃薯根际可培养细菌和放线菌数量明显减少，真菌数量显著增加，细菌与真菌数量的比值（B/F值）随连作年限的增加而减小，从轮作L0的114.22×10~3降低到连作5年L5的1.08×10~3，马铃薯连作使根际微生物群落组成由“细菌”型向“真菌”型转变。
DGGE指纹图谱结果显示，马铃薯连作使根际土壤中细菌和真菌的种群结构均发生显著变化：马铃薯连作使根际土壤中细菌优势种群的个体数明显减少，其中对照处理L0条带数量最多，有68个操作分类单元（OTU），其它处理的条带数量均少于L0，特别是连作3年以后，条带数量减少的更明显，连作3-5年分别比对照（轮作）处理减少了19.12%、41.18%和29.41%；真菌优势种群的个体数明显增多，连作1-5年（L1-L5）处理的操作分类单元（OTU）分别比对照处理增加了38%、38%、31%、46%和77%。由此可见，马铃薯连作造成根际土壤微生物的选择性适应，出现某些种群富集，而某些种群数量降低的现象。聚类分析结果表明，连作马铃薯根际土壤细菌和真菌种群结构的相似性在相邻的连作年限间都比较高，然而随着连作年限的增加，各处理的相似性越来越低。
从真菌DGGE条带的测序结果来看，随着连作年限的增加，马铃薯根际土壤土传病害病原菌Fusarium oxysporum和Fusarium solani数量明显增加，而Chaetomium globosum作为一种生防菌，至连作5年，数量明显减少，连作使根际土壤中土传病害病原菌种群过渡成为优势微生物种群。
马铃薯连作使根系总吸收面积和活跃吸收面积均明显降低，与对照L0相比，在现蕾期，L2和L4根系总吸收面积分别下降了26.07%和51.18%，根系活跃吸收面积分别下降了17.14%和48.57%；在块茎膨大期，L2和L4处理根系总吸收面积分别下降了27.10%和80.06%，根系活跃吸收面积分别下降了32.68%和85.37%，且处理间均表现出显著差异。根系活力也表现出降低的趋势，在现蕾期，根系活力大小表现为L2＞L0＞L4，且处理间差异显著；在块茎膨大期则表现为L0＞L2＞L4，与L0相比，L2和L4处理根系活力分别下降了5.25%和69.29%，说明在轻度连作的情况下，马铃薯根系的活力并未表现出显著性的降低，甚至在营养生长时期有所上升。由此可见，连作造成了马铃薯根际微生态系统失衡，导致根际微生态环境恶化。
2.连作对马铃薯植株的生物效应研究
马铃薯叶片SOD、POD和CAT活性都随着生育期的推进而升高。在现蕾期，随着连作年限的增加，叶片SOD和POD活性随之升高，CAT活性先升高后降低，但三者的连作处理均高于对照；至块茎膨大期，叶片SOD和CAT活性都表现为：连作2年>连作4年>对照，各处理间差异不显著，POD活性先降低后升高，但连作处理均显著低于对照。叶片中MDA含量随着随生长发育的推进而升高，并且随连作年限的增加而增加，叶片细胞质膜的过氧化作用加重。
马铃薯连作造成植株生长发育受阻，外观表现为植株矮小，单株主茎数减少，各器官生物量均随着连作年限的增加而降低，对单株结薯数影响不大，但是却显著降低了植株的单株结薯重量，最终表现为块茎产量大幅度降低。
综上所述，马铃薯连作使根际微生物群落组成及种群结构发生明显变化，特别是根际土壤中土传病害病原菌大量增加，生防菌数量减少，根系活力和吸收面积显著降低，造成根际微生态系统失衡，根际微生态环境恶化，进而影响植株正常的代谢活动，使马铃薯生长发育受阻，最终导致块茎产量大幅度下降。
Long-time continuous cropping potato has been one of the big problems thatrestrict the yield and serious pests and diseases, a field experiment was carried out inJingtai Tiaoshan Farm in Gansu Province in2010, the rotation cropping field is CK(continuous cropping potato for zero years, L0), the rest is the continuous croppingpotato from one year to five years (L1-L5), starting from the point of view of therhizosphere micro-ecological, using the PCR-DGGE molecular fingerprintingtechniques to study rhizosphere microbial community composition, bacteria andfungal population structure changes and root physiological activity change are thecharacteristic rhizosphere micro-ecological environment change, and after therhizosphere micro-ecological environment change, the leaf protective enzymesactivity and malondialdehyde content, the biological traits of plants and yield in thecondition of continuous cropping potato. A preliminary study on the effect and themechanism of potato continuous cropping obstacles to provide a theoretical basis toreduce the potato cropping obstacles. The main results were as follows:
1. Effects of potato continuous cropping on the rhizosphere micro-ecological
The number of culturable rhizosphere microorganisms had significantlydifferent response for potato continuous cropping. bacterial response was sensitive,actinomycetes response was a little lag. With the continuous cropping yearsincreasing, the number of bacteria and actinomycetes decreased in the potatorhizosphere soil, while the number of fungi increased significantly; The ratio ofbacteria and fungi (B/F-value) decreased with the continuous cropping yearsincreasing, from rotation cropping L0for114.22×10~3to continuous cropping potatofor5years L5for1.08×10~3, potato continuous cropping made the rhizosphere fromthe “bacterial” type to “fungi” type.
The results for DGGE were as follows, continuous cropping potato maderhizosphere bacteria and fungi population structure change significantly: continuouscropping potato significantly reduced the numbers of individuals of the bacteriadominant populations in rhizosphere soil，the numbers of visible bands were the most in L0, which were68the operational taxonomic unit (OTU), other treatmentswere much less than L0, the numbers of visible bands were significantly reducedafter continuous cropping three years, compared with CK, continuous cropping three,four and five years were respectively reduced19.12%,41.18%and29.41%.Continuous cropping significantly increased the numbers of individuals of the fungidominant populations, compared with CK, the OTU of continuous croppingtreatments from one to five years were increased by38%,38%,31%,46%and77%respectively. All the above illustrated that continuous cropping potato causedselective adaptation of rhizosphere soil microorganisms, which enriched somepopulations or reduced certain populations. The cluster analysis showed that, thesimilarity of bacterial and fungal population structure were higher in the adjacentcontinuous cropping years, with the continuous cropping years increasing, however,the similarity of each treatment is becoming less and less.
From the sequencing results of the fungal DGGE bands, with the increase ofcontinuous cropping years, the number of the potato rhizosphere soil-bornepathogens Fusarium oxysporum and Fusarium solani increased significantly, whileChaetomium globosum, as a biocontrol species, was reduced significantly incontinuous cropping five years. Potato continuous cropping caused pathogen fungalpopulations become the dominant microbial populations in rhizosphere.
The total absorbing area and actively absorbing area of roots were significantlyreduced by the continuous cropping potato. Compared with L0, the total absorbingarea of roots of L2and L4were respectively reduced26.07%and51.18%, and theactively absorbing area of roots were respectively reduced17.14%and48.57%inthe budding stage. The total absorbing area of roots were respectively reduced27.10%and80.06%, the actively absorbing area of roots were respectively reduced32.68%and85.37%in the tuber expansion stage, and the difference was significantbetween treatments. And root vigor displayed a decreasing trend, in the buddingstage, the results are as follows: L2＞L0＞L4, and the difference was significantbetween treatments. In the tuber expansion stage, the results are as follows: L0＞L2 ＞L4, compared with L0, L2and L4were respectively reduced5.25%and69.29%.Therefore, when the continuous cropping was mild, root vigor of potato had notsignificantly been reduced, even which was increased in the nutritional growthperiod.
2. Effects of potato continuous cropping on the plant biological roles
Potato leaf SOD, POD and CAT activity were increased with the advance of thegrowth period. In the budding stage, SOD and POD activity were both increasedwith the continuous cropping years increasing, CAT activity first was increased andthen was decreased. To the tuber enlargement stage, leaf SOD and CAT activitieswere as follows: continuous cropping for three years＞continuous cropping for fiveyears＞CK, and the difference was not significant between treatments, POD activityof continuous cropping treatments were less than CK. MDA content was increasedwith the advance of the growth period, and which was also increased with thecontinuous cropping years increasing, and the cell membrane peroxidation effect ofthe leaf was increased.
Growth of potato was restraibed under the condition of continuous cropping,such as plant short, Per plant main stems were reduced, the biomass of each organwere decreased with the continuous cropping years increasing, The tuber number ofper plant was little affected, however, per plant tuber weight was significantlyreduced, the ultimate expression was that the tuber yield had a significant reduction.
In conclusion, the potato rhizosphere microorganisms community compositionand population structure had a change significantly under the condition ofcontinuous cropping, the number of the potato rhizosphere soil-borne diseasepathogens was significantly increased, but the number of biocontrol strain wasreduced obviously, the rhizosphere micro-ecological environment destruction, whichhindered the normal metabolic activity of plant system, the growth of potato wasrestrained eventually, the ultimate result was that the tuber yield had a significantreduction.
1.连作对马铃薯根际微生态环境的影响
马铃薯根际可培养微生物的数量对连作的反应程度明显不同，细菌对连作的反应较敏感，放线菌反应稍滞后。随着连作年限的增加，马铃薯根际可培养细菌和放线菌数量明显减少，真菌数量显著增加，细菌与真菌数量的比值（B/F值）随连作年限的增加而减小，从轮作L0的114.22×10~3降低到连作5年L5的1.08×10~3，马铃薯连作使根际微生物群落组成由“细菌”型向“真菌”型转变。
DGGE指纹图谱结果显示，马铃薯连作使根际土壤中细菌和真菌的种群结构均发生显著变化：马铃薯连作使根际土壤中细菌优势种群的个体数明显减少，其中对照处理L0条带数量最多，有68个操作分类单元（OTU），其它处理的条带数量均少于L0，特别是连作3年以后，条带数量减少的更明显，连作3-5年分别比对照（轮作）处理减少了19.12%、41.18%和29.41%；真菌优势种群的个体数明显增多，连作1-5年（L1-L5）处理的操作分类单元（OTU）分别比对照处理增加了38%、38%、31%、46%和77%。由此可见，马铃薯连作造成根际土壤微生物的选择性适应，出现某些种群富集，而某些种群数量降低的现象。聚类分析结果表明，连作马铃薯根际土壤细菌和真菌种群结构的相似性在相邻的连作年限间都比较高，然而随着连作年限的增加，各处理的相似性越来越低。
从真菌DGGE条带的测序结果来看，随着连作年限的增加，马铃薯根际土壤土传病害病原菌Fusarium oxysporum和Fusarium solani数量明显增加，而Chaetomium globosum作为一种生防菌，至连作5年，数量明显减少，连作使根际土壤中土传病害病原菌种群过渡成为优势微生物种群。
马铃薯连作使根系总吸收面积和活跃吸收面积均明显降低，与对照L0相比，在现蕾期，L2和L4根系总吸收面积分别下降了26.07%和51.18%，根系活跃吸收面积分别下降了17.14%和48.57%；在块茎膨大期，L2和L4处理根系总吸收面积分别下降了27.10%和80.06%，根系活跃吸收面积分别下降了32.68%和85.37%，且处理间均表现出显著差异。根系活力也表现出降低的趋势，在现蕾期，根系活力大小表现为L2＞L0＞L4，且处理间差异显著；在块茎膨大期则表现为L0＞L2＞L4，与L0相比，L2和L4处理根系活力分别下降了5.25%和69.29%，说明在轻度连作的情况下，马铃薯根系的活力并未表现出显著性的降低，甚至在营养生长时期有所上升。由此可见，连作造成了马铃薯根际微生态系统失衡，导致根际微生态环境恶化。
2.连作对马铃薯植株的生物效应研究
马铃薯叶片SOD、POD和CAT活性都随着生育期的推进而升高。在现蕾期，随着连作年限的增加，叶片SOD和POD活性随之升高，CAT活性先升高后降低，但三者的连作处理均高于对照；至块茎膨大期，叶片SOD和CAT活性都表现为：连作2年>连作4年>对照，各处理间差异不显著，POD活性先降低后升高，但连作处理均显著低于对照。叶片中MDA含量随着随生长发育的推进而升高，并且随连作年限的增加而增加，叶片细胞质膜的过氧化作用加重。
马铃薯连作造成植株生长发育受阻，外观表现为植株矮小，单株主茎数减少，各器官生物量均随着连作年限的增加而降低，对单株结薯数影响不大，但是却显著降低了植株的单株结薯重量，最终表现为块茎产量大幅度降低。
综上所述，马铃薯连作使根际微生物群落组成及种群结构发生明显变化，特别是根际土壤中土传病害病原菌大量增加，生防菌数量减少，根系活力和吸收面积显著降低，造成根际微生态系统失衡，根际微生态环境恶化，进而影响植株正常的代谢活动，使马铃薯生长发育受阻，最终导致块茎产量大幅度下降。
Long-time continuous cropping potato has been one of the big problems thatrestrict the yield and serious pests and diseases, a field experiment was carried out inJingtai Tiaoshan Farm in Gansu Province in2010, the rotation cropping field is CK(continuous cropping potato for zero years, L0), the rest is the continuous croppingpotato from one year to five years (L1-L5), starting from the point of view of therhizosphere micro-ecological, using the PCR-DGGE molecular fingerprintingtechniques to study rhizosphere microbial community composition, bacteria andfungal population structure changes and root physiological activity change are thecharacteristic rhizosphere micro-ecological environment change, and after therhizosphere micro-ecological environment change, the leaf protective enzymesactivity and malondialdehyde content, the biological traits of plants and yield in thecondition of continuous cropping potato. A preliminary study on the effect and themechanism of potato continuous cropping obstacles to provide a theoretical basis toreduce the potato cropping obstacles. The main results were as follows:
1. Effects of potato continuous cropping on the rhizosphere micro-ecological
The number of culturable rhizosphere microorganisms had significantlydifferent response for potato continuous cropping. bacterial response was sensitive,actinomycetes response was a little lag. With the continuous cropping yearsincreasing, the number of bacteria and actinomycetes decreased in the potatorhizosphere soil, while the number of fungi increased significantly; The ratio ofbacteria and fungi (B/F-value) decreased with the continuous cropping yearsincreasing, from rotation cropping L0for114.22×10~3to continuous cropping potatofor5years L5for1.08×10~3, potato continuous cropping made the rhizosphere fromthe “bacterial” type to “fungi” type.
The results for DGGE were as follows, continuous cropping potato maderhizosphere bacteria and fungi population structure change significantly: continuouscropping potato significantly reduced the numbers of individuals of the bacteriadominant populations in rhizosphere soil，the numbers of visible bands were the most in L0, which were68the operational taxonomic unit (OTU), other treatmentswere much less than L0, the numbers of visible bands were significantly reducedafter continuous cropping three years, compared with CK, continuous cropping three,four and five years were respectively reduced19.12%,41.18%and29.41%.Continuous cropping significantly increased the numbers of individuals of the fungidominant populations, compared with CK, the OTU of continuous croppingtreatments from one to five years were increased by38%,38%,31%,46%and77%respectively. All the above illustrated that continuous cropping potato causedselective adaptation of rhizosphere soil microorganisms, which enriched somepopulations or reduced certain populations. The cluster analysis showed that, thesimilarity of bacterial and fungal population structure were higher in the adjacentcontinuous cropping years, with the continuous cropping years increasing, however,the similarity of each treatment is becoming less and less.
From the sequencing results of the fungal DGGE bands, with the increase ofcontinuous cropping years, the number of the potato rhizosphere soil-bornepathogens Fusarium oxysporum and Fusarium solani increased significantly, whileChaetomium globosum, as a biocontrol species, was reduced significantly incontinuous cropping five years. Potato continuous cropping caused pathogen fungalpopulations become the dominant microbial populations in rhizosphere.
The total absorbing area and actively absorbing area of roots were significantlyreduced by the continuous cropping potato. Compared with L0, the total absorbingarea of roots of L2and L4were respectively reduced26.07%and51.18%, and theactively absorbing area of roots were respectively reduced17.14%and48.57%inthe budding stage. The total absorbing area of roots were respectively reduced27.10%and80.06%, the actively absorbing area of roots were respectively reduced32.68%and85.37%in the tuber expansion stage, and the difference was significantbetween treatments. And root vigor displayed a decreasing trend, in the buddingstage, the results are as follows: L2＞L0＞L4, and the difference was significantbetween treatments. In the tuber expansion stage, the results are as follows: L0＞L2 ＞L4, compared with L0, L2and L4were respectively reduced5.25%and69.29%.Therefore, when the continuous cropping was mild, root vigor of potato had notsignificantly been reduced, even which was increased in the nutritional growthperiod.
2. Effects of potato continuous cropping on the plant biological roles
Potato leaf SOD, POD and CAT activity were increased with the advance of thegrowth period. In the budding stage, SOD and POD activity were both increasedwith the continuous cropping years increasing, CAT activity first was increased andthen was decreased. To the tuber enlargement stage, leaf SOD and CAT activitieswere as follows: continuous cropping for three years＞continuous cropping for fiveyears＞CK, and the difference was not significant between treatments, POD activityof continuous cropping treatments were less than CK. MDA content was increasedwith the advance of the growth period, and which was also increased with thecontinuous cropping years increasing, and the cell membrane peroxidation effect ofthe leaf was increased.
Growth of potato was restraibed under the condition of continuous cropping,such as plant short, Per plant main stems were reduced, the biomass of each organwere decreased with the continuous cropping years increasing, The tuber number ofper plant was little affected, however, per plant tuber weight was significantlyreduced, the ultimate expression was that the tuber yield had a significant reduction.
In conclusion, the potato rhizosphere microorganisms community compositionand population structure had a change significantly under the condition ofcontinuous cropping, the number of the potato rhizosphere soil-borne diseasepathogens was significantly increased, but the number of biocontrol strain wasreduced obviously, the rhizosphere micro-ecological environment destruction, whichhindered the normal metabolic activity of plant system, the growth of potato wasrestrained eventually, the ultimate result was that the tuber yield had a significantreduction.
引文
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