Using competitive and facilitative interactions in intercropping systems enhances crop productivity and nutrient-use efficiency

This paper reviews recent research on the processes involved in the yield advantage in wheat (Triticum aestivum L.)/maize (Zea mays L.), wheat/soybean [Glycine max (L.) Merr.], faba bean (Vicia faba L.)/maize, peanut (Arachis hypogaea L.)/maize and water convolvulus (Ipomoea aquatica Forsk.)/maize intercropping. In wheat/maize and wheat/soybean intercropping systems, a significant yield increase of intercropped wheat over sole wheat was observed, which resulted from positive effects of the border row and inner rows of intercropped wheat. The border row effect was due to interspecific competition for nutrients as wheat had a higher competitive ability than either maize or soybean had. There was also compensatory growth, or a recovery process, of subordinate species such as maize and soybean, offsetting the impairment of early growth of the subordinate species. Finally, both dominant and subordinate species in intercropping obtain higher yields than that in corresponding sole wheat, maize or soybean. We summarized these processes as the `competition-recovery production principle'. We observed interspecific facilitation, where maize improves iron nutrition in intercropped peanut, faba bean enhances nitrogen and phosphorus uptake by intercropped maize, and chickpea facilitates P uptake by associated wheat from phytate-P. Furthermore, intercropping reduced the nitrate content in the soil profile as intercropping uses soil nutrients more efficiently than sole cropping.     

Effects of rhizobial inoculation, cropping systems and growth stages on endophytic bacterial community of soybean roots

Interspecific interactions and soil nitrogen supply levels affect intercropping productivity. We hypothesized that interspecific competition can be alleviated by increasing N application rate and yield advantage can be obtained in competitive systems. A field experiment was conducted in Wuwei, Gansu province in 2007 and 2008 to study intercropping of faba bean/maize, wheat/maize, barley/maize and the corresponding monocultures of faba bean (Vicia faba L.), wheat (Triticum aestivum L.), barley (Hordeum vulgare L.) and maize (Zea mays L.) with N application rates of 0, 75, 150, 225 and 300 kg N ha^?1. Total land equivalent ratios (TLER) were 1.22 for faba bean/maize, 1.16 for wheat/maize, and 1.13 for barley/maize intercropping over the 2-year study period. Maize was overyielding when intercropped with faba bean, but underyielding when intercropped with wheat or barley according to partial land equivalent ratios (PLER) based on grain yields of individual crops in intercropping and sole cropping. There was an interspecific facilitation between intercropped faba bean and maize, and interspecific competition between maize and either wheat or barley. The underyielding of maize was higher when intercropped with barley than with wheat. Fertilizer N alleviated competitive interactions in intercrops with adequate fertilizer N at 225 kg ha^?1. Yield advantage of intercropping can be acquired with adequate nitrogen supply, even in an intensive competitive system such as barley/maize intercropping. This is important when using intercropping to develop intensive farming systems with high inputs and high outputs.     

种间互作和施氮对蚕豆/玉米间作生态系统地上部和地下部生长的影响

为河西走廊绿洲灌区豆科/禾本科间作体系的养分管理提供科学依据,于2007年在武威绿洲农业试验站应用田间原位根系行分隔技术研究了蚕豆/玉米种间互作和施氮对玉米抽雄期的根系空间分布、根系形态和作物地上部生长的影响。研究结果表明:种间互作和施氮均增加了玉米和蚕豆在纵向和横向两个尺度上的根重密度、根长密度、根表面积、根系体积。根长密度和根表面积与两种作物产量和氮素吸收均呈正相关,而与蚕豆的根瘤重呈负相关;抽雄期的土壤含水量与玉米产量和养分吸收呈显著的负相关。玉米根系可以占据蚕豆地下部空间,但蚕豆的根却较少到间作玉米的地下部空间,也就是间作后增加了玉米根系水平尺度的生态位。蚕豆和玉米根系主要分布分别在0-40 cm浅土层和0-60 cm 土层,且间作玉米根系在60-120 cm比单作和分隔的多。因此,种间互作和施氮扩大了两作物根系纵向和横向的空间生态位,改变了作物根系形态,即扩展了两者水分和养分吸收的生态位,增加了作物吸收养分的有效空间,从而提高了间作生态系统的生产力。     

Wheat and faba bean intercropping changes phenolic acids from roots to rhizosphere

The changed phenolic acids (PAs) allelochemicals exuded by the roots induced by interspecific interactions is related to intercropping alleviates soil-borne disease. However, the presence of PAs in roots and root exudations and their rhizodeposition under intercropping are still unclear. Hydroponic and soil experiments of wheat, faba bean, and wheat intercropped with faba bean were conducted, and the major compositions and contents of PAs in roots, root exudations, and rhizospheric soil were determined. The results showed that ρ-hydroxybenzoic, vanillic, and syringic acids were the major components of PAs in roots, root exudations, and rhizospheric soil in a wheat and faba bean intercropping system. The compositions and percentages of PAs in roots of faba bean were altered when faba bean intercropped with wheat. The total exudation rate of PAs in root exudations was decreased by 30%–60% under the wheat and faba bean intercropping (W//F) system as compared to mono-cropped faba bean (MF). ρ-hydroxybenzoic acid was identified in the root exudation of both MF and mono-cropped wheat (MW), but not detected in the intercropping on 60 days after transplanting. Vanillic acid was only detected in the root exudation of MF on 30 days after transplanting. The rhizodepostion of vanillic and cumaric acid were decreased at both branching and pod setting stages in W//F as compared to MF. In conclusion, interspecific interaction changed the compositions and contents of PAs in faba bean roots and root exudations. W//F constrained vanillic acid exuded by roots and decreased vanillic and coumaric acid rhizodeposition by faba bean, which provides insight into root-soil interactions in the intercropping systems.     

An overview of rhizosphere processes related with plant nutrition in major cropping systems in China

Rhizosphere processes of individual plants have been widely investigated since 1904 when the term “rhizosphere” was first put forward. However, little attention has been paid to rhizosphere effects at an agro-ecosystem level. This paper presents recent research on the rhizosphere processes in relation to plant nutrition in main cropping systems in China. In the peanut (Arachis hypogaea L.)/maize (Zea mays L.) intercropping system, maize was found to improve the Fe nutrition of peanut through influencing its rhizosphere processes, suggesting an important role of phytosiderophores released from Fe-deficient maize. Intercropping between maize and faba bean (Vicia faba L.) was found to improve nitrogen and phosphorus uptake in the two crops compared with corresponding sole crop. There was a higher land equivalent ratio (LER) in the intercropping system of maize and faba bean than the treatment of no root interactions between the two crops. The increased yield of maize intercropped with faba bean resulted from an interspecific facilitation in nutrient uptake, depending on interspecific root interactions of the two crops. In the rotation system of rice (Oryza sativa L.)-wheat (Triticum aestivum L.) crops, Mn deficiency in wheat was caused by excessive Mn uptake by rice and Mn leaching from topsoil to subsoil due to periodic cycles of flooding and drying. However, wheat genotypes tolerant to Mn deficiency tended to distribute more roots to deeper soil layer and thus expand their rhizosphere zones in the Mn-deficient soils and utilize Mn from the subsoil. Deep ploughing also helped root penetration into subsoil and was propitious to correcting Mn deficiency in wheat rotated with rice. In comparison, oilseed rape (Brassica napus L.) took up more Mn than wheat through mobilizing sparingly soluble soil Mn due to acidification and reduction processes in the rhizosphere. Thus, oilseed rape was tolerant to the Mn-deficient conditions in the rice-oilseed rape rotation. Oxidation reactions on root surface of rice also resulted in the formation of Fe plaque in the rice rhizosphere. Large amounts of Zn were accumulated on the Fe plaque. Zinc uptake by rice plants increased as Fe plaque formed, but decreased at high amounts of Fe plaque. It is suggested that to fine-tune cropping patterns and optimize nutrient management based on a better understanding of rhizosphere processes at an agro-ecosystem level is crucial for increasing nutrient use efficiency and developing sustainable agriculture in China.     

玉米、小麦和大麦与蚕豆间作体系不同根系分隔方式对蚕豆结瘤的影响

本实验选取3种对土壤氮素竞争能力不同的禾本科作物大麦(Hordeum vulgare)、小麦(Triticum aestivum)和玉米(Zea mays)分别与蚕豆(Vicia faba)间作, 建立对土壤氮素竞争能力不同的作物组合; 并采用3种分隔方式(塑料膜分隔、尼龙网分隔和无分隔)建立同一作物组合条件下作物种间根系相互作用的不同强度, 来研究不同作物组合及种间根系相互作用强度对蚕豆结瘤的影响。结果如下: (1)蚕豆的结瘤并未随3种禾本科作物氮素竞争能力的增强而增加, 但是3种间作体系蚕豆的结瘤却均表现出无分隔处理多于塑料膜分隔处理, 即同一间作体系种间根系相互作用越强, 越有利于蚕豆结瘤的产生, 存在种间互利作用; (2)在玉米/蚕豆间作体系中, 无分隔处理的蚕豆根瘤数目和根瘤重显著高于塑料膜分隔处理, 分别高出67.5%和70.1%; 在大麦/蚕豆间作体系中也表现出无分隔处理的根瘤重显著高于塑料膜分隔处理(高出46.3%); (3)玉米/蚕豆间作体系与小麦/蚕豆和大麦/蚕豆间作体系相比, 无分隔处理时土壤氮素含量显著高于后2个间作体系, 但是玉米/蚕豆间作体系对蚕豆结瘤的促进作用更强。上述结果表明, 在蚕豆/玉米间作体系中, 玉米促进蚕豆生物固氮除了氮素竞争机制外, 还可能存在其它机制。     

Nitrogen Fixation of Faba Bean (Vicia faba L.) Interacting with a Non-legume in Two Contrasting Intercropping Systems

A field experiment was carried out to quantify biological nitrogen fixation (BNF) using the ¹⁵N isotope natural abundance method in maize (Zea mays L.)/faba bean (Vicia faba L.) and wheat (Triticum aestivum L.)/faba bean intercropping systems. Faba bean was yielding more in the maize/faba bean intercropping, but not in the wheat/faba bean intercropping. Biomass, grain yield and N acquisition of faba bean were significantly increased when intercropped with maize, and decreased significantly with wheat, irrespective of N-fertilizer application, indicating that the legume could gain or lose productivity in an intercropping situation. There was yield advantage of maize/faba bean intercropping, but no in wheat/faba bean intercropping. The grain yield of the faba bean intercropped with maize was greater than that of faba bean monoculture due to increases of the stems per plant and the pods per stem of faba bean. N fertilization inhibited N fixation of faba bean in maize/faba bean and wheat/faba bean intercropping and faba bean monoculture. The responses of different cropping systems to N-fertilizer application, however, were not identical, with competitive intercropping (wheat/faba bean) being more sensitive than facilitative intercropping (maize/faba bean). Intercropping increased the percentage of N derived from air (%Ndfa) of the wheat/faba bean system, but not that of the maize/faba bean system when no N fertilizer was applied. When receiving 120 kg N/ha, however, intercropping did not significantly increase %Ndfa either in the wheat/faba bean system or in the maize/faba bean system in comparison with faba bean in monoculture. The amount of shoot N derived from air (Ndfa), however, increased significantly when intercropped with maize, irrespective of N-fertilizer application. Ndfa decreased when intercropped with wheat, albeit not significantly at 120 kg N/ha. Ndfa was correlated more closely with dry matter yield, grain yield and competitive ratio, than with %Ndfa. This indicates that that total dry matter yield (sink strength), not %Ndfa, was more critical for the legume to increase Ndfa. The results suggested that N fixation could be improved by yield maximization in an intercropping system.     

Interspecific complementary and competitive interactions between intercropped maize and faba bean

Interspecific complementary and competitive interactions between maize (Zea mays L. cv. Zhongdan No. 2) and faba bean (Vicia faba L. cv. Linxia Dacaidou) in maize/faba bean intercropping systems were assessed in two field experiments in Gansu province, northwestern China, plus a microplot experiment in one treatment of one of the field experiments in which root system partitions were used to determine interspecific root interactions. Intercropping effects were detected, with land equivalent ratio values of 1.21–1.23 based on total (grain+straw) yield and 1.13–1.34 based on grain yield. When two rows of maize were intercropped with two rows of faba bean, both total yield and grain yield of both crop species were significantly higher than those of sole maize and faba bean on an equivalent area basis. When two rows of pea (Pisum sativum L. cv. Beijing No. 5) were intercropped with two rows of faba bean, neither total yield nor grain yield of faba bean was higher than of sole faba bean on an equivalent area basis. Interspecific competition between maize and faba bean was relatively weak, with mean relative crowding coefficients of 0.99–1.02 for maize and 1.55–1.59 for faba bean. The microplot experiment in which partitions were placed between root systems showed a significant positive yield effect on maize when the root systems intermingled freely (no partition) or partly (400 mesh nylon net partition) compared with no interspecific root interaction (plastic sheet partition). This revised version was published online in June 2006 with corrections to the Cover Date.     

The role of maize root size in phosphorus uptake and productivity of maize/faba bean and maize/wheat intercropping systems

Interspecific root/rhizosphere interactions affect phosphorus (P) uptake and the productivity of maize/faba bean and maize/wheat intercropping systems. The aim of these experiments was to determine whether manipulation of maize root growth could improve the productivity of the two intercropping systems. Two near isogenic maize hybrids (the larger-rooted T149 and smaller-rooted T222) were intercropped with faba bean and wheat, under conditions of high- and low-P availability. The larger-rooted T149 showed greater competitive ability than the smaller-rooted T222 in both maize/faba bean and maize/wheat intercropping systems. The higher competitive ability of T149 improved the productivity of the maize/faba bean intercropping system in P-sufficient conditions. In maize/wheat intercropping systems, root growth, shoot biomass, and P uptake of maize were inhibited by wheat, regardless of the P-supply. Compared with T222, the larger-rooted T149 suffered less in the intercropping systems. The total biomass of the maize/wheat intercropping system was higher for wheat/T149 than for wheat/T222 under low-P conditions. These data suggested that genetic improvement of maize root size could enhance maize growth and its ability to compete for P resources in maize/faba bean and maize/wheat intercropping systems. In addition, depending on the P availability, larger maize roots could increase the productivity of intercropping systems.     

The Dynamic Process of Interspecific Interactions of Competitive Nitrogen Capture between Intercropped Wheat (Triticum aestivum L.) and Faba Bean (Vicia faba L.)

Wheat (Triticum aestivum L.)/faba bean (Vicia faba L.) intercropping shows significant overyielding and high nitrogen (N)-use efficiency, but the dynamics of plant interactions have rarely been estimated. The objective of the present study was to investigate the temporal dynamics of competitive N acquisition between intercropped wheat and faba bean with the logistic model. Wheat and faba bean were grown together or alone with limited N supply in pots. Data of shoot and root biomass and N content measured from 14 samplings were fitted to logistic models to determine instantaneous rates of growth and N uptake. The superiority of instantaneous biomass production and N uptake shifted from faba bean to wheat with their growth. Moreover, the shift of superiority on N uptake occurred 7–12 days earlier than that of biomass production. Interspecific competition stimulated intercropped wheat to have a much earlier and stronger superiority on instantaneous N uptake compared with isolated wheat. The modeling methodology characterized the temporal dynamics of biomass production and N uptake of intercropped wheat and faba bean in different planting systems, which helps to understand the underlying process of plant interaction for intercropping plants.     

供肥对小麦间作蚕豆群体产量及根系的调控

利用田间,池栽,盆栽等方法,对不同N,P水平下小麦间作蚕豆群体根系的时空分布及产量效应进行了研究。结果表明,供P对小麦-蚕豆间作系统有明显的增产作用,与不供P相比,田间试验中供P处理间作小麦和蚕豆分别提高了48.39%和16.69%,池栽试验中群体产量增产幅度为20.07%-43.14%。间作蚕豆经济产量增幅为58.46%-78.78%,小麦间作蚕豆多作系统2种作物根密度生长高峰期交错出现,小麦的峰值出现早于蚕豆,其中,小麦和蚕豆根干重最大值分别出现在抽穗期和成熟期,而根长的最大值分别出现在拔节期和成熟期,这在一定程度上减轻了共生期2种作物对水肥的竞争,也是此群体增产增效的原因之一,施P处理间作小麦的根重,根长和根表面积分别比不施P处理提高54.33%,48.88%和47.00%;施N处理间作小麦的根重,根长和根表面积分别比不施N的增加15.25%,11.61%和11.46%。间作小麦根重的57.61%和蚕豆的69.20%分布在0-30cm土层,随施P水平的提高,总根干重及根密度都趋于增加,且深层土壤中根系增加明显。     

Crop nitrogen use and soil mineral nitrogen accumulation under different crop combinations and patterns of strip intercropping in northwest China

Increasing crop nitrogen use efficiency while also simultaneously decreasing nitrogen accumulation in the soil would be key steps in controlling nitrogen pollution from agricultural systems. Long-term field experiments were started in 2003 to study the effects of intercropping on crop N use and soil mineral N accumulation in wheat (Triticum aestivum L. cv 2014)/maize (Zea mays L. cv Shendan16), wheat/faba bean (Vicia faba L. cv Lincan No. 5) and maize/faba bean intercropping and monocropping systems. Monocropping was compared with two types of strip intercropping: continuous intercropping (two crops intercropped continuously on the same strips of land every year) and rotational intercropping (two crops grown adjacently and rotated to the other crop??s strip every year). Maize/faba bean intercropping had greater crop N uptake than did wheat/faba bean or wheat/maize. Wheat/maize accumulated more mineral N in the top 140 cm of the soil profile during the co-growth stage from maize emergence to maturity of wheat or faba bean. Continuously intercropped maize substantially decreased soil mineral N accumulation under wheat and faba bean rows (60?C100 cm soil depth) at maize harvest. Soil mineral N accumulation under wheat rows increased with rotational intercropping with faba bean. Rotational intercropping may potentially alleviate the adverse effects of wheat on N use by other crops and increase the nitrogen harvest index of wheat, maize and faba bean. Intercropping using species with different maturity dates may be more effective in increasing crop N use efficiency and decreasing soil mineral N accumulation.     

Changes in soil phosphorus fractions following sole cropped and intercropped maize and faba bean grown on calcareous soil

Aims

This study aimed to investigate the effects of coexistence with faba bean, a phosphorus (P)-efficient crop, on soil-accumulated P use by a maize/faba bean intercropping system on dynamic changes in soil P pool.

Methods

Maize and faba bean were grown in P-accumulated soil as either sole cropping or intercropping. After one year (Stage I) or four years (Stage II) of no P application, soil samples were collected respectively and analyzed for soil P pools using sequential fractionation. Aboveground biomass and P content were annually measured from 2013 to 2016 to assess the annual P balance.

Results

The intercropped maize/faba bean system showed a P-uptake advantage, with a Land Equivalent Ratio (LER) ranging from 1.2 to 1.5. The average shoot P content over the four years in intercropped maize and faba bean was significantly greater than that of the corresponding sole crops by 29% and 30%, respectively. Over the three-year P depletion period, the three cropping systems primarily depleted the 1 M HCl-P_i fraction, followed by sole maize, which depleted the NaOH-P_i and concentrated HCl-P_o fractions. Sole faba bean depleted the alkali-soluble P_o fraction (extracted by NaHCO₃ and NaOH), and the intercropped maize/faba bean system depleted the conc. HCl-P_o fraction, which was similar to the effect of sole maize.

Conclusions

Both sole crops and intercrops mainly depleted 1 M HCl-P_i, but differed in P_o depletion. Sole maize and maize/faba bean intercropping depleted the sparingly labile P_o fraction, while sole faba bean depleted the labile and moderately labile P_o fractions.

     

大豆和蚕豆苗期根系生长特征的比较

大豆和蚕豆是西北地区间套作生产中广泛种植的豆科作物,但二者在与禾本科作物间作时差异十分明显,蚕豆相对于大豆具有更强的竞争能力.从大豆和蚕豆的根系入手,研究了大豆和蚕豆在苗期根系生长发育和形态的差异.采用PVC管砂培装置,通过扫描仪扫描根系,用图像分析软件WinRHIZO进行了研究.结果表明,出苗42 d时,蚕豆根表面积是大豆根表面积的2.61倍;出苗14 d时,大豆根系在整个砂层中均有分布,而蚕豆根系只分布在表层;大豆根较细,大部分根的直径为0.2～1.0 mm,相对于大豆而言,蚕豆根较粗,大部分根直径为0.5～1.5 mm.蚕豆根较粗,所以有更大的表面积,而且它的根系分布在上层的比例大于大豆,这些因素决定了它的吸收能力强于大豆,能更好地利用上层土壤养分.这是蚕豆竞争能力强于大豆的原因之一.     

小麦/蚕豆间作作物生长曲线的模拟及种间互作分析

物种间的相互作用与间作产量优势的形成密切相关,但很少有人注意到种间互作动态.本研究通过2年田间定位试验,运用Logistic分析模拟了不同种植模式(小麦单作、蚕豆单作和小麦/蚕豆间作)和不同磷水平下[P₀,施磷量(P₂O₅)为0 kg·hm^-2(对照);P₁,施磷量(P₂O₅)为45 kg·hm^-2;P₂,施磷量(P₂O₅)为90 kg·hm^-2]单间作小麦、蚕豆的生长模型,分析了作物种间互作的动态变化.结果表明: 小麦/蚕豆间作使小麦产量提高了10.5%~18.6%,蚕豆产量却降低了4.8%~12.3%,但间作系统仍具有产量优势,土地当量比(LER)和相对拥挤系数(K)分别为1.01~1.15 和1.12~3.20.小麦和蚕豆的产量及关键生长参数均受磷水平调控,但LER和K并不受磷水平影响.与单作相比,间作小麦的最大生长速率(R_max)和最初生长速率(r)分别提高21.8%~38.7%和20.7%~38.9%,但间作对蚕豆的关键生长参数无影响.在小麦、蚕豆的生长初期,不同磷水平下,单间作作物的生长曲线无差异;间作群体以种间竞争为主,无间作生物量优势(LER<1,K<1).当蚕豆达到最大生长速率(T_max)后,间作显著提高了小麦的生长速率,降低了小麦的种内竞争压力,表现出间作生物量和产量优势(LER>1,K>1).总之,在不同的生长发育阶段,小麦、蚕豆的相互作用不同,间作提高了中后期小麦的生长速率,为间作优势的形成奠定了基础.     

小麦与蚕豆间作体系根系形态与磷吸收的定量解析

豆科与禾本科作物间作能够改变作物根系生长,但不同施磷水平下间作-根系形态-磷吸收之间的关系尚未明确。本研究通过田间定位试验和根箱模拟试验,研究不同种植模式(小麦单作、蚕豆单作和小麦-蚕豆间作)和不同磷水平下小麦和蚕豆的产量、生物量、磷吸收及根系形态特征,分析探讨不同施磷条件下小麦-蚕豆间作对根系形态和磷吸收的影响。结果表明: 根箱试验中,与单作相比,间作小麦的根干重、根冠比分别增加21.2%、61.5%,地上部干重降低14.6%,根系磷含量和总吸磷量分别提高23.8%和12.1%;间作蚕豆的地上部干重、根干重、根冠比、总根长和根体积分别增加16.5%、47.3%、24.0%、3.5%和8.4%,间作蚕豆地上部磷含量、根系磷含量和总吸磷量分别提高14.7%、26.2%和21.5%。田间试验中,与单作相比,分蘖期间作小麦地上部磷吸收量降低8.7%,而在拔节期、抽穗期、灌浆期和成熟期分别提高40.6%、19.7%、7.8%和12.4%;但种间互作导致开花期、结荚期和成熟期间作蚕豆的地上部磷吸收量分别降低9.8%、9.0%和5.2%。偏最小二乘法(PLS)回归分析表明,小麦的根表面积和根体积、蚕豆的根表面积对作物磷吸收的贡献最大,在低磷胁迫条件下,间作同时提高了小麦和蚕豆的根体积和根表面积,促进了磷的吸收。总之,在缺磷或低磷条件下,种间互作扩大了小麦和蚕豆根土接触面积,促进了苗期磷的吸收,为后期间作优势的形成奠定了基础。     

小麦蚕豆间作对蚕豆根际微生物群落功能多样性的影响及其与蚕豆枯萎病发生的关系

通过田间小区试验,研究了小麦与蚕豆间作对蚕豆枯萎病发生和根际微生物代谢功能多样性的影响。结果表明,小麦与蚕豆间作使蚕豆枯萎病的发病率和病情指数分别比单作显著降低20%和30.4%。与单作处理相比,间作显著增加了蚕豆和小麦根际微生物对31种碳源的平均利用率（AWCD）,其中间作蚕豆的AWCD值最高,比单作增加82.7%,单作蚕豆最低。间作蚕豆和间作小麦根际微生物的Shannon多样性指数与丰富度指数均显著高于单作,间作使蚕豆和小麦的丰富度指数分别增加29.2%和30.3%。根际微生物对六类碳源的利用强度百分比以糖类、羧酸类和氨基酸最高,分别为41.96%,19.80%和18.13%。主成分分析表明,小麦与蚕豆间作改变了根际微生物的群落组成;相关分析表明,糖类、羧酸类和氨基酸类碳源是区分单间作处理差异的主要碳源,其中氨基酸类碳源是最敏感的碳源。小麦与蚕豆间作增加了根际微生物活性,提高了Shannon多样性指数和丰富度指数,改变了微生物群落功能多样性,是抑制蚕豆枯萎病的有效措施。该研究为阐明根际微生物功能多样性变化在间作体系病害控制中的作用与机制奠定了理论基础。     

间作减轻蚕豆枯萎病的微生物和生理机制

通过田间小区试验,研究了小麦与蚕豆间作对蚕豆枯萎病发病率、病情指数、根际镰刀菌数量、蚕豆根系抗氧化酶活性和膜质过氧化的影响.采用Biolog ECO板分析了根际土壤微生物的代谢功能多样性,通过高效液相色谱(HPLC)测定了蚕豆根际土壤中酚酸含量.结果表明: 与单作蚕豆相比,小麦与蚕豆间作有降低蚕豆枯萎病发病率的趋势;发病盛期和发病末期,间作使蚕豆枯萎病的病情指数比单作显著降低15.8%和22.8%,明显提高了蚕豆根际微生物活性(AWCD值),根际微生物的Shannon多样性指数显著提高4.4%和5.3%,丰富度指数显著提高19.4%和37.1%.主成分分析表明,发病盛期和发病末期,间作明显改变了蚕豆根际微生物的群落结构,蚕豆根际镰刀菌数量分别降低53.8%和33.1%;并显著降低了蚕豆根际土壤中对羟基苯甲酸、香草酸、丁香酸、阿魏酸、苯甲酸和肉桂酸的含量.发病盛期和发病末期,间作蚕豆根系的过氧化物酶(POD)和过氧化氢酶(CAT)活性分别显著提高了20.0%、31.3%和38.5%、66.7%,丙二醛(MDA)含量显著降低36.3%和46.3%;发病初期间作对蚕豆根系的POD、CAT酶活性和MDA含量无显著影响.小麦与蚕豆间作显著提高了蚕豆根际微生物的活性、多样性和根系抗氧化酶活性,降低了蚕豆根际土壤中酚酸含量和膜质过氧化程度,减少了土壤镰刀菌的数量,从而提高了蚕豆对枯萎病的抗性,降低了枯萎病的危害程度.     

不同品种小麦与蚕豆间作对蚕豆枯萎病的防治及其机理

通过田间小区试验和水培试验,研究了3个品种小麦（云麦42、云麦47和绵阳29）与蚕豆间作对蚕豆地上部生物量、枯萎病发生、根际微生物代谢功能多样性和枯萎病病原菌尖孢镰刀菌数量的影响,分析了3个不同小麦品种低分子量根系分泌物组分（糖、氨基酸和有机酸）的差异.结果表明： 云麦42与蚕豆间作（YM42/B）和云麦47与蚕豆间作（YM47/B）使蚕豆地上部生物量分别增加16.6%和13.4%,使蚕豆枯萎病病情指数分别降低47.6%和23.3%,绵阳29与蚕豆间作（MY29/B）对蚕豆地上部生物量和枯萎病病情指数均无显著影响.与单作蚕豆相比,YM42/B和YM47/B处理均显著提高了蚕豆根际微生物的平均颜色变化率（AWCD)及对碳源的总利用强度和Biolog ECO板中各种碳源的利用,明显改变了蚕豆根际微生物群落结构,并显著降低了蚕豆根际的尖孢镰刀菌数量;MY29/B处理对AWCD值、碳源总利用强度和Biolog ECO板中各种碳源的利用及蚕豆根际尖孢镰刀菌数量均无显著影响,也未明显改变蚕豆根际的微生物群落结构.3个小麦品种根系分泌物中可溶性糖、游离氨基酸总量和有机酸总量均表现为MY29＞YM47＞YM42.MY29根系分泌物中丝氨酸、谷氨酸、甘氨酸、缬氨酸、蛋氨酸、苯丙氨酸、赖氨酸的含量均显著高于YM42和YM47;精氨酸为YM42、YM47所特有,而亮氨酸为MY29所特有.MY29和YM47根系分泌物中检出酒石酸、苹果酸、柠檬酸、琥珀酸、延胡索酸和顺乌头酸6种有机酸,而YM42根系分泌物中检出洒石酸、苹果酸、柠檬酸和延胡索酸4种有机酸,且MY29和YM47根系分泌物中苹果酸含量显著高于YM42.小麦与蚕豆间作提高了蚕豆根际微生物的活性和碳源利用强度,改变了根际微生物的群落结构,降低了蚕豆根际尖孢镰刀菌的数量,促进了蚕豆生长,抑制了蚕豆枯萎病的发生,但小麦与蚕豆间作控病效果受小麦品种的影响,表明间作系统中非寄主作物根系分泌物的差异是影响间作对土传病害控制的重要因素.     

Mixed cropping with maize combined with moderate UV-B radiations lead to enhanced flavonoid production and root growth in faba bean

Abstract

Flavonoids have recently been proposed to function as developmental regulators and/or signaling molecules under biotic or abiotic stress. The aim of this study was to determine the composition and concentration of fiavonoid aglycones (kaempferol, luteolin, and quercetin) in faba bean shoots and roots, as affected by interspecific root interactions with maize and moderate UV-B radiation. Independent of the UV-B treatment, interspecific root interactions with maize enhanced the concentration of both quercetin and luteolin in the faba bean shoots and roots by 50 and 97.8%, respectively, and improved the root length by 14.6%. In addition, moderate UV-B radiation facilitated a systematic increase of both aglycones in both shoots and roots without affecting plant growth. To our knowledge, this report is the first work documenting the response of faba bean flavonoids to interspecific root interactions with maize and moderate UV-B radiation, and it provides a new perspective for understanding interspecific interactions.