小豆连作障碍中自毒机理研究
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摘要
小豆生育期短，耐瘠、耐阴，适应性广，是我国主要的食用豆类作物之一，也是我国重要的出口农产品和优势农产品，其总产量和出口量均居世界第一位。连作障碍是制约小豆产品质量和产量的主要限制因素。据调查，连作种植小豆发病率一般为30%，严重田块高达80%。因此，揭示小豆自毒作用与连作障碍的之间关系，研究自毒作用下小豆植株生长、土壤环境和产量的响应机制，对提高我国小豆产量和品质，促进小豆产业可持续发展具有重要意义。
论文以小豆高产品种冀红9218为材料，通过田间试验和室内分析相结合，鉴定分析小豆各器官中主要自毒物质的种类和成分；在人工模拟条件下，研究了主效自毒物质在土壤环境条件下对小豆植株活性氧代谢，土壤微生物、养分和酶活性变化、矿质营养及光合作用的作用影响。得出如下主要结论：
（1）小豆的根、茎、叶及根际土壤浸提液对种子萌发和幼苗生长抑制作用明显，且随浸提液浓度的增大而抑制作用逐渐增强。而且不同浸提液对小豆种子萌发指标影响效果存在差异，以叶片浸提液的抑制作用最为明显；不同器官浸提液对小豆幼苗生长的影响也有相似的变化趋势，其抑制效应由强到弱依次为枯叶、鲜叶、土壤、根系、茎秆。
（2）利用GC-MS技术分别对小豆根、茎、叶和根际土壤浸提液中的自毒物质成分和含量进行鉴定，结果表明，不同器官浸提液中自毒物质的成分和含量之间存在明显不同，叶片浸提液中共检测到55种化学物质，小豆根系、根际土壤浸提液和茎秆分别检测到化感成分40、43、33种。研究发现，不同器官浸提液中含量较高的化感物质是邻苯二甲酸、肉桂酸和对羟基苯甲酸。通过生物测试验证，这3种酚酸类物质均具有自毒作用，对小豆幼苗根系的生长有明显抑制作用，且有浓度梯度效应，其中以邻苯二甲酸自毒作用最明显，其次是肉桂酸，以对羟基苯甲酸对发芽率、胚根生长和植株干重的抑制作用最弱。
（3）盆栽试验表明，不同浓度的肉桂酸和邻苯二甲酸均能抑制小豆根系生长，并随浓度的增加抑制作用增强。而且以邻苯二甲酸的抑制作用更为明显。邻苯二甲酸浓度在10mmol.L-1浓度下对根干重、根长、根体积、根表面积、根系活力、根瘤数和根瘤鲜重均产生显著抑制作用。此外，肉桂酸处理后小豆单株粒重分别较对照降低了1.1，2.1，3.0g，邻苯二甲酸较对照下降了2.6、3.2和4.3g。与对照相比，百粒重分别较对照减少1.4、2.1、3.5g，2.3、3.2、4.1g。两种自毒物质对产量的影响，以邻苯二甲酸抑制效果明显。
（4）肉桂酸和邻苯二甲酸降低了小豆植株叶片SOD和CAT活性，随着处理时间的增加，SOD和CAT活性前期升高，后期下降。在第28d时，邻苯二甲酸处理下，SOD活性下降幅度最大，各浓度处理分别下降了9.01%，18.45%，35.41%；CAT活性下降了14.59%，23.33%和31.57%。但是，随着处理浓度和胁迫时间的增加，POD活性和MDA含量呈增加的趋势。
（5）肉桂酸和邻苯二甲酸对小豆叶片光合能力有明显的抑制效应。整个生育过程，不同处理之间，小豆叶片光合指标（叶绿素含量，可溶性蛋白，净光合速率，气孔导度，蒸腾速率）表现出先上升后下降的变化趋势，处理初期差异不显著，随着生育期的推进和自毒物质的积累，不同处理之间，与对照相比，差异达显著水平。但是胞间CO2浓度始终与对照差异不显著。相同处理时间下，不同浓度与抑制效果呈正相关关系。两种自毒物质对光合能力的抑制作用，以邻苯二甲酸的作用更强。
（6）肉桂酸和邻苯二甲酸均使土壤酶活性和微生物数量发生改变。随着自毒物质不断积累，土壤中过氧化氢酶、脲酶、碱性磷酸酶和转化酶活性逐渐下降；以转化酶为例，在同一浓度处理下，随肉桂酸胁迫时间的延长，转化酶活性在第14-21d达到最大值，而后下降幅度明显，到胁迫后期(42d)时，不同浓度处理下，转化酶活性分别比对照下降了18.22%、29.35%和38.49%。对邻苯二甲酸处理对根际土壤中转化酶的活性也表现出相同的变化趋势，分别显著下降了23.48%，34.85%和49.71%。在自毒物质作用下，根际土壤中细菌、真菌、放线菌数量所占比例发生改变。土壤中的细菌、放线菌的数量明显减少，而真菌数量显著增加。
（7）与对照相比，肉桂酸和邻苯二甲酸抑制小豆根际土壤中养分含量，降低全P、K向速效P、K元素的转化。尤其是在10mmol.L-1浓度下，两种自毒物质均显著抑制了对土壤中全P，K的吸收，但是对氮素的影响差异不明显。
（8）随着干旱程度的增加，不同叶片浸提液浓度下，小豆根系活力呈现明显下降过程，二者交互作用对小豆根系活力表现为抑制效应。而且重度干旱胁迫后，叶片浸提液浓度0.04和0.16g·ml-1处理与对照相比差异达到极显著水平。营养亏缺条件下，小豆叶片叶绿素含量和根系活力分别在0.04和0.16g·ml-1叶片浸提液处理下与对照间差异达到显著水平。小豆根系MDA含量随着浸提液浓度增大而逐渐升高，表现出促进丙二醛的积累。二者相比，以缺K条件下抑制效果更明显。
Adzuki bean (Vigna angularis) with a short growth days, shade plant, resistance-acidityand good adaptability is a important bean crop in China, meanwhile it is also importantagricultural product in export, and yield and export amount is maximum all over the world.Replant problem was a key factor for restraining yield and quality in adzuki bean. Replant canincrease rate of disease infecting, which can reach by30%-80%. Therefore, there was animportant meaning for overcoming replant problem based on studying relationship betweenautotoxin and replant problem in Adzuki bean.
Jihong9218was used as a material and analyzed the kinds and composition of autotoxinin different organs of adzuki bean by yield test and lab. Under manual simulation, we focusedon researching effect of reactive oxygen metabolism, soil microorganism, nutrition andenzyme variation, mineral nutrition and photosynthesis. The main results were as follows:
(1)The leach liquor of different organs and root area in adzuki bean has negative effecton seed sprouting and seedling growth, and this effect was increasing with increasing leachliquor density. Autotoxin was site difference during seed sprouting in adzuki bean, and therewas significant autotoxin in leaf leach liquor. The order of negative effect in different organswas as follows: Flag> fresh leaf> soil> root> stem.
(2) Different organs and soil leach liquor in root were measured by using GC-MSmethod. The results showed that: the kinds and content of allelochemicals had large differencein leaf, stem, root, root area. There were55,43,40,33kinds of chemical materials in leachliquor of leaf, root, stem, root area, respectively. Different organs and soil leach liquor hadsame allelochemicals, and they were phthalic acid, cinnamic acid, p-hydroxybenzoic acid,respectively. However, these allelochemicals had different content in different organs.According to bioassay, these materials had inhibitional effect for root growth in adzuki bean,and this inhibitional effect was increasing with increasing density. The order of negative effectwas as follows: phthalic acid> cinnamic acid> p-hydroxybenzoic acid.
(3) Phthalic acid and cinnamic acid can restrain root growth, and this effect was increasewith increasing density, meanwhile phthalic acid had better negative effect than cinnamic acid.Phthalic acid had significant negative effect in dry weight of root, root length, root volume, root area, root vitality, the number of root nodule and root fresh weight under10m mol.l-1. Inadditional, abortive grain,100-seed weight and the pod number per plant had negative effectwith increasing density of phthalic acid and cinnamic acid. Compared with CK, grain weightper plant of adzuki bean was decrease by1.1g,2.1g,3.0g and2.6g,3.2g,4.3g by cinnamicacid processing and Phthalic acid processing, respectively. This result showed that phthalicacid had better effect on yield than cinnamic acid.
(4) Phthalic acid and cinnamic acid can reduce activity of SOD and CAT. Activity ofSOD and CAT was increase in early stage and decreasing in later stage with increasing inprocessing time. In28days after seedling, activity of SOD and CAT were decrease by9.01%，18.45%，35.41%and14.59%，23.33%,31.57%by phthalic acid processing,respectively. Activity of POD and content of MDA were increase with increasing inprocessing density and stress time.
(5) Phthalic acid and cinnamic acid had negative effect on content of chlorophyll, netphotosynthetic rate, stomatal conductance and transpiration rate, meanwhile they had doublenegative effect on intercellular CO2density, but there was no significant difference indifferent treatments. Under0.1m mol.L-1density, content of chlorophyll, net photosyntheticrate, stomatal conductance and transpiration rate were increase firstly and decrease latterlywith increasing stress time. Under10m mol.L-1density, content of chlorophyll, netphotosynthetic rate, stomatal conductance and transpiration rate were decrease gradually withincreasing stress time.
(6) Compared with CK, the treatment of two autotoxins can decrease number ofbacterias, actinomyces and microorganisms in rhizosphere soil and increase number offunguses, meanwhile every treatment can decrease activity of CAT, protease, phosphatase andsucrase in rhizosphere soil. Invertin was taken as example, under the same density, activity ofinvertin was maximum in14-21d then decrease apparently with stress time prolong bycinnamic acid. In42days, activity of invertin was decrease by18.22%、29.35%,38.49%compared with CK in different density processing. There was the same trend by phthalic acidprocessing, and that decreased by23.48%，34.85%,49.71%, respectively. Under autotoxinprocessing, the number of bacterium, fungus, and actinomyces was vary in soil. The numberof bacterium and actinomyces were decrease, and the number of fungus was increase.
(7) Phthalic acid and cinnamic acid can restrain nutrition to be absorbed in root ofadzuki bean. Under10m mol.L-1density, absorbing of P and K were restrained by twoautotoxins, which decreased use efficiency of P, K and had no significant effect on N.
(8)Activity of root in adzuki bean was decrease in leaf leach liquor with drought stressincreasing, and drought and leach liquor had significant negative effect on activity of root in adzuki bean. Under high drought stress, there was significant difference (P<0.01) than CK intreatments of0.04g ml-1,0.16g ml-1in leaf leach liquor. Under nutrition stress, the content ofchlorophyll and activity of root in adzuki bean was significant difference (P<0.05) than CKunder treatments of0.04g ml-1,0.16g ml-1in leaf leach liquor. The content of MDA wasincrease with leach liquor density increasing, and MDA accumulated gradually. Comparedleach liquor with drought stress, there was more negative effect in K deficiency.
论文以小豆高产品种冀红9218为材料，通过田间试验和室内分析相结合，鉴定分析小豆各器官中主要自毒物质的种类和成分；在人工模拟条件下，研究了主效自毒物质在土壤环境条件下对小豆植株活性氧代谢，土壤微生物、养分和酶活性变化、矿质营养及光合作用的作用影响。得出如下主要结论：
（1）小豆的根、茎、叶及根际土壤浸提液对种子萌发和幼苗生长抑制作用明显，且随浸提液浓度的增大而抑制作用逐渐增强。而且不同浸提液对小豆种子萌发指标影响效果存在差异，以叶片浸提液的抑制作用最为明显；不同器官浸提液对小豆幼苗生长的影响也有相似的变化趋势，其抑制效应由强到弱依次为枯叶、鲜叶、土壤、根系、茎秆。
（2）利用GC-MS技术分别对小豆根、茎、叶和根际土壤浸提液中的自毒物质成分和含量进行鉴定，结果表明，不同器官浸提液中自毒物质的成分和含量之间存在明显不同，叶片浸提液中共检测到55种化学物质，小豆根系、根际土壤浸提液和茎秆分别检测到化感成分40、43、33种。研究发现，不同器官浸提液中含量较高的化感物质是邻苯二甲酸、肉桂酸和对羟基苯甲酸。通过生物测试验证，这3种酚酸类物质均具有自毒作用，对小豆幼苗根系的生长有明显抑制作用，且有浓度梯度效应，其中以邻苯二甲酸自毒作用最明显，其次是肉桂酸，以对羟基苯甲酸对发芽率、胚根生长和植株干重的抑制作用最弱。
（3）盆栽试验表明，不同浓度的肉桂酸和邻苯二甲酸均能抑制小豆根系生长，并随浓度的增加抑制作用增强。而且以邻苯二甲酸的抑制作用更为明显。邻苯二甲酸浓度在10mmol.L-1浓度下对根干重、根长、根体积、根表面积、根系活力、根瘤数和根瘤鲜重均产生显著抑制作用。此外，肉桂酸处理后小豆单株粒重分别较对照降低了1.1，2.1，3.0g，邻苯二甲酸较对照下降了2.6、3.2和4.3g。与对照相比，百粒重分别较对照减少1.4、2.1、3.5g，2.3、3.2、4.1g。两种自毒物质对产量的影响，以邻苯二甲酸抑制效果明显。
（4）肉桂酸和邻苯二甲酸降低了小豆植株叶片SOD和CAT活性，随着处理时间的增加，SOD和CAT活性前期升高，后期下降。在第28d时，邻苯二甲酸处理下，SOD活性下降幅度最大，各浓度处理分别下降了9.01%，18.45%，35.41%；CAT活性下降了14.59%，23.33%和31.57%。但是，随着处理浓度和胁迫时间的增加，POD活性和MDA含量呈增加的趋势。
（5）肉桂酸和邻苯二甲酸对小豆叶片光合能力有明显的抑制效应。整个生育过程，不同处理之间，小豆叶片光合指标（叶绿素含量，可溶性蛋白，净光合速率，气孔导度，蒸腾速率）表现出先上升后下降的变化趋势，处理初期差异不显著，随着生育期的推进和自毒物质的积累，不同处理之间，与对照相比，差异达显著水平。但是胞间CO2浓度始终与对照差异不显著。相同处理时间下，不同浓度与抑制效果呈正相关关系。两种自毒物质对光合能力的抑制作用，以邻苯二甲酸的作用更强。
（6）肉桂酸和邻苯二甲酸均使土壤酶活性和微生物数量发生改变。随着自毒物质不断积累，土壤中过氧化氢酶、脲酶、碱性磷酸酶和转化酶活性逐渐下降；以转化酶为例，在同一浓度处理下，随肉桂酸胁迫时间的延长，转化酶活性在第14-21d达到最大值，而后下降幅度明显，到胁迫后期(42d)时，不同浓度处理下，转化酶活性分别比对照下降了18.22%、29.35%和38.49%。对邻苯二甲酸处理对根际土壤中转化酶的活性也表现出相同的变化趋势，分别显著下降了23.48%，34.85%和49.71%。在自毒物质作用下，根际土壤中细菌、真菌、放线菌数量所占比例发生改变。土壤中的细菌、放线菌的数量明显减少，而真菌数量显著增加。
（7）与对照相比，肉桂酸和邻苯二甲酸抑制小豆根际土壤中养分含量，降低全P、K向速效P、K元素的转化。尤其是在10mmol.L-1浓度下，两种自毒物质均显著抑制了对土壤中全P，K的吸收，但是对氮素的影响差异不明显。
（8）随着干旱程度的增加，不同叶片浸提液浓度下，小豆根系活力呈现明显下降过程，二者交互作用对小豆根系活力表现为抑制效应。而且重度干旱胁迫后，叶片浸提液浓度0.04和0.16g·ml-1处理与对照相比差异达到极显著水平。营养亏缺条件下，小豆叶片叶绿素含量和根系活力分别在0.04和0.16g·ml-1叶片浸提液处理下与对照间差异达到显著水平。小豆根系MDA含量随着浸提液浓度增大而逐渐升高，表现出促进丙二醛的积累。二者相比，以缺K条件下抑制效果更明显。
Adzuki bean (Vigna angularis) with a short growth days, shade plant, resistance-acidityand good adaptability is a important bean crop in China, meanwhile it is also importantagricultural product in export, and yield and export amount is maximum all over the world.Replant problem was a key factor for restraining yield and quality in adzuki bean. Replant canincrease rate of disease infecting, which can reach by30%-80%. Therefore, there was animportant meaning for overcoming replant problem based on studying relationship betweenautotoxin and replant problem in Adzuki bean.
Jihong9218was used as a material and analyzed the kinds and composition of autotoxinin different organs of adzuki bean by yield test and lab. Under manual simulation, we focusedon researching effect of reactive oxygen metabolism, soil microorganism, nutrition andenzyme variation, mineral nutrition and photosynthesis. The main results were as follows:
(1)The leach liquor of different organs and root area in adzuki bean has negative effecton seed sprouting and seedling growth, and this effect was increasing with increasing leachliquor density. Autotoxin was site difference during seed sprouting in adzuki bean, and therewas significant autotoxin in leaf leach liquor. The order of negative effect in different organswas as follows: Flag> fresh leaf> soil> root> stem.
(2) Different organs and soil leach liquor in root were measured by using GC-MSmethod. The results showed that: the kinds and content of allelochemicals had large differencein leaf, stem, root, root area. There were55,43,40,33kinds of chemical materials in leachliquor of leaf, root, stem, root area, respectively. Different organs and soil leach liquor hadsame allelochemicals, and they were phthalic acid, cinnamic acid, p-hydroxybenzoic acid,respectively. However, these allelochemicals had different content in different organs.According to bioassay, these materials had inhibitional effect for root growth in adzuki bean,and this inhibitional effect was increasing with increasing density. The order of negative effectwas as follows: phthalic acid> cinnamic acid> p-hydroxybenzoic acid.
(3) Phthalic acid and cinnamic acid can restrain root growth, and this effect was increasewith increasing density, meanwhile phthalic acid had better negative effect than cinnamic acid.Phthalic acid had significant negative effect in dry weight of root, root length, root volume, root area, root vitality, the number of root nodule and root fresh weight under10m mol.l-1. Inadditional, abortive grain,100-seed weight and the pod number per plant had negative effectwith increasing density of phthalic acid and cinnamic acid. Compared with CK, grain weightper plant of adzuki bean was decrease by1.1g,2.1g,3.0g and2.6g,3.2g,4.3g by cinnamicacid processing and Phthalic acid processing, respectively. This result showed that phthalicacid had better effect on yield than cinnamic acid.
(4) Phthalic acid and cinnamic acid can reduce activity of SOD and CAT. Activity ofSOD and CAT was increase in early stage and decreasing in later stage with increasing inprocessing time. In28days after seedling, activity of SOD and CAT were decrease by9.01%，18.45%，35.41%and14.59%，23.33%,31.57%by phthalic acid processing,respectively. Activity of POD and content of MDA were increase with increasing inprocessing density and stress time.
(5) Phthalic acid and cinnamic acid had negative effect on content of chlorophyll, netphotosynthetic rate, stomatal conductance and transpiration rate, meanwhile they had doublenegative effect on intercellular CO2density, but there was no significant difference indifferent treatments. Under0.1m mol.L-1density, content of chlorophyll, net photosyntheticrate, stomatal conductance and transpiration rate were increase firstly and decrease latterlywith increasing stress time. Under10m mol.L-1density, content of chlorophyll, netphotosynthetic rate, stomatal conductance and transpiration rate were decrease gradually withincreasing stress time.
(6) Compared with CK, the treatment of two autotoxins can decrease number ofbacterias, actinomyces and microorganisms in rhizosphere soil and increase number offunguses, meanwhile every treatment can decrease activity of CAT, protease, phosphatase andsucrase in rhizosphere soil. Invertin was taken as example, under the same density, activity ofinvertin was maximum in14-21d then decrease apparently with stress time prolong bycinnamic acid. In42days, activity of invertin was decrease by18.22%、29.35%,38.49%compared with CK in different density processing. There was the same trend by phthalic acidprocessing, and that decreased by23.48%，34.85%,49.71%, respectively. Under autotoxinprocessing, the number of bacterium, fungus, and actinomyces was vary in soil. The numberof bacterium and actinomyces were decrease, and the number of fungus was increase.
(7) Phthalic acid and cinnamic acid can restrain nutrition to be absorbed in root ofadzuki bean. Under10m mol.L-1density, absorbing of P and K were restrained by twoautotoxins, which decreased use efficiency of P, K and had no significant effect on N.
(8)Activity of root in adzuki bean was decrease in leaf leach liquor with drought stressincreasing, and drought and leach liquor had significant negative effect on activity of root in adzuki bean. Under high drought stress, there was significant difference (P<0.01) than CK intreatments of0.04g ml-1,0.16g ml-1in leaf leach liquor. Under nutrition stress, the content ofchlorophyll and activity of root in adzuki bean was significant difference (P<0.05) than CKunder treatments of0.04g ml-1,0.16g ml-1in leaf leach liquor. The content of MDA wasincrease with leach liquor density increasing, and MDA accumulated gradually. Comparedleach liquor with drought stress, there was more negative effect in K deficiency.
引文
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