Root Interactions in a Maize/Soybean Intercropping System Control Soybean Soil-Borne Disease,Red Crown Rot

Background

Within-field multiple crop species intercropping is well documented and used for disease control, but the underlying mechanisms are still unclear. As roots are the primary organ for perceiving signals in the soil from neighboring plants, root behavior may play an important role in soil-borne disease control.

Principal Findings

In two years of field experiments, maize/soybean intercropping suppressed the occurrence of soybean red crown rot, a severe soil-borne disease caused by Cylindrocladium parasiticum (C. parasiticum). The suppressive effects decreased with increasing distance between intercropped plants under both low P and high P supply, suggesting that root interactions play a significant role independent of nutrient status. Further detailed quantitative studies revealed that the diversity and intensity of root interactions altered the expression of important soybean PR genes, as well as, the activity of corresponding enzymes in both P treatments. Furthermore, 5 phenolic acids were detected in root exudates of maize/soybean intercropped plants. Among these phenolic acids, cinnamic acid was released in significantly greater concentrations when intercropped maize with soybean compared to either crop grown in monoculture, and this spike in cinnamic acid was found dramatically constrain C. parasiticum growth in vitro.

Conclusions

To the best of our knowledge, this study is the first report to demonstrate that intercropping with maize can promote resistance in soybean to red crown rot in a root-dependent manner. This supports the point that intercropping may be an efficient ecological strategy to control soil-borne plant disease and should be incorporated in sustainable agricultural management practices.     

玉米/大豆间作条件下的作物根系生长及水分吸收

通过田间试验研究了玉米/大豆条带间作群体的根系分布及土壤水分吸收规律.结果表明：水分充足条件下,土壤剖面内玉米和大豆根系的分布模式近似于三角形;玉米根系水平分布范围较大,侧向伸展长度约为58 cm,16～22 cm土层的玉米根系侧向伸展最远,玉米根系不仅分布于间作条带行间,而且生长到大豆条带的行间;大豆根系水平分布于相对有限的区域内,侧向伸展长度约为26 cm.作物根质量密度随着距作物行（玉米或大豆）距离的增加而减少,玉米行和边行大豆根质量密度的90%分布于0～30 cm土层.距玉米行10 cm处玉米的根质量密度高于大豆,距玉米行20 cm处大豆的根质量密度大于玉米,两种作物根质量密度的85%都分布于0～30 cm土层内.间作条带内水分变化主要集中在0～30 cm土层,水分变化量依次为：玉米区域>大豆区域>条带行间.表明在水分充足条件下,间作作物优先在自己的区域吸水,根系混合区吸水滞后发生.     

Plant-Plant-Microbe Mechanisms Involved in Soil-Borne Disease Suppression on a Maize and Pepper Intercropping System

Background

Intercropping systems could increase crop diversity and avoid vulnerability to biotic stresses. Most studies have shown that intercropping can provide relief to crops against wind-dispersed pathogens. However, there was limited data on how the practice of intercropping help crops against soil-borne Phytophthora disease.

Principal Findings

Compared to pepper monoculture, a large scale intercropping study of maize grown between pepper rows reduced disease levels of the soil-borne pepper Phytophthora blight. These reduced disease levels of Phytophthora in the intercropping system were correlated with the ability of maize plants to form a “root wall” that restricted the movement of Phytophthora capsici across rows. Experimentally, it was found that maize roots attracted the zoospores of P. capsici and then inhibited their growth. When maize plants were grown in close proximity to each other, the roots produced and secreted larger quantities of 2,4-dihydroxy-7-methoxy-2H-1,4-benzoxazin-3(4H)-one (DIMBOA) and 6-methoxy-2-benzoxazolinone (MBOA). Furthermore, MBOA, benzothiazole (BZO), and 2-(methylthio)-benzothiazole (MBZO) were identified in root exudates of maize and showed antimicrobial activity against P. capsici.

Conclusions

Maize could form a “root wall” to restrict the spread of P. capsici across rows in maize and pepper intercropping systems. Antimicrobe compounds secreted by maize root were one of the factors that resulted in the inhibition of P. capsici. These results provide new insights into plant-plant-microbe mechanisms involved in intercropping systems.     

Phosphorus benefits from grain-legume crops to subsequent maize grown on acid soils of southern Cameroon

We conducted field experiments over 2 years on two acid soils of southern Cameroon to test whether efficient uptake and use of phosphorus (P) from less available sources by grain legume genotypes could benefit subsequent rotational maize. We grew two crops each year. For the first crop we grew 4 genotypes of soybean and of cowpea, plus maize. For the second crop we grew maize. The first crops were fertilized with 0, 90 kg P ha⁻¹ as phosphate rock (PR) or 30 kg P ha⁻¹ as triple super phosphate (TSP). P application highly significantly increased shoot dry matter, P uptake, N₂ fixation and grain yields of the grain legumes with TSP generally more effective than PR. Two of the soybean and two of the cowpea genotypes were more efficient at using P. Only the P-efficient soybean and cowpea genotypes increased subsequent maize yields. Yields of the subsequent maize grown in rotation were significantly correlated with shoot P uptake for which the quantity of P applied with the crop residues of the pre-crop appeared to be a major factor. We also grew the grain legumes in nutrient solutions and measured organic acid-anion exudation from roots, root-surface phosphatase-activity, and root morphological characteristics. Enhanced exudation of organic acid anions from roots of P-deprived plants might have contributed to the P acquisition efficiency under field conditions of the P-efficient cowpea genotypes and one of the P-efficient soybean genotypes. A higher activity of root-surface acid phosphatase might have been important for the other P-efficient soybean genotype. The results show, that the potential positive rotational effect of cowpea and soybean on the acid, highly P-sorbing soils of southern Cameroon depends on breeding and using P-efficient genotypes when sparingly soluble and suboptimal rates of soluble P fertilizers are used. Section Editor: N. J. Barrow     

间作对植株生长及养分吸收和根际环境的影响

通过盆栽实验研究了线辣椒和玉米间作对其植株生长、矿质养分吸收、根际环境以及铁载体分泌的影响,以探索间作促进铁、磷等养分吸收利用的可能生理机制.结果表明:(1)与单作相比,间作线辣椒地上部干重降低23.0%,根系干重增加44.2%,玉米地上部和根系的干重分别增加8.7%和22.9%;间作线辣椒根冠比和根系活力分别显著提高86.4%和29.8%;间作线辣椒、玉米叶绿素含量分别显著提高12.6%和7.8%.(2)与单作相比,间作线辣椒的铁、锌、锰含量分别增加1.50倍、1.39倍和1.34%,而间作玉米则无显著变化;间作线辣椒和玉米的钙含量都显著低于相应单作,氮含量没有显著变化,但磷、钾含量显著增加.(3)间作线辣椒和玉米的根际土、非根际土的酸性磷酸酶活性及根系酸性磷酸酶活性都显著高于相应单作,而其根际土和非根际土的pH值无显著变化;间作玉米根系的铁载体分泌比单作减少32.8%,间作线辣椒根系的铁还原酶活性是单作的1.10倍.研究发现,线辣椒/玉米间作能通过影响根际生物学特征和化学过程提高植株的铁、锌、磷和钾养分水平,缓解养分胁迫,是一种很有推广价值的种植模式.     

Nutrient transfer between the root zones of soybean and maize plants connected by a common mycorrhizal mycelium

The objective of the study was to determine whether nutrient fluxes mediated by hyphae of vesicular-arbuscular mycorrhizal (VAM) fungi between the root zones of grass and legume plants differ with the legume's mode of N nutrition. The plants, nodulating or nonnodulating isolines of soybean [ Glycine max (L.) Merr.], were grown in association with a dwarf maize ( Zea mays L.) cultivar in containers which interposed a 6-cm-wide root-free soil bridge between legume and grass container compartments. The bridge was delimited by screens (44 μm) which permitted the passage of hyphae, but not of roots and minimized non VAM interactions between the plants. All plants were colonized by the VAM fungus Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe. The effects of N input to N-sufficient soybean plants through N₂-fixation or N-fertilization on associated maize-plant growth and nutrition were compared to those of an N-deficient (nonnodulating, unfertilized) soybean control. Maize, when associated with the N-fertilized soybean, increased 19% in biomass, 67% in N content and 77% in leaf N concentration relative to the maize plants of the N-deficient association. When maize was grown with nodulated soybean, maize N content increased by 22%, biomass did not change, but P content declined by 16%. Spore production by the VAM fungus was greatest in the soils of both plants of the N-fertilized treatment. The patterns of N and P distribution, as well as those of the other essential elements, indicated that association with the N-fertilized soybean plants was more advantageous to maize than was association with the N₂-fixing ones.     

Phosphate absorption by air-stressed root systems

Summary Root systems from plants grown in nutrient solution were exposed to air and either transferred to fresh nutrient solution containing ³²P-labeled phosphate or placed in a psychrometer to determine their water potential. The amount of ³²P absorbed by maize and soybean roots in the hour following their exposure to air was proportional to their water potential at the time they were transferred. Some cells, probably located in the stele, were more resistant to moisture stress than others. Absorption of ³²P by all cells was severely inhibited by water potentials below-12 to-15 bars. Nearly normal amounts of the radioisotope and total phosphate were absorbed within 72 hr following root exposure of 4 of 5 species of detopped plants; some phosphorus was lost to the nutrient solution. Uptake of ³²P by passive processes was increased slightly by exposure of roots of intact maize plants to air, but the increase did not compensate for the substantial reduction in actively-absorbed ³²P.     

Peanut/maize intercropping induced changes in rhizosphere and nutrient concentrations in shoots.

A greenhouse study was conducted to investigate the rhizosphere effects on iron (Fe), phosphorus (P), nitrogen (N), potassium (K), calcium (Ca), zinc (Zn), and manganese (Mn) nutrition in peanut plants (Arachis hypogaea L.) by intercropping them with maize (Zea mays L.). In addition, we studied the release of phytosiderophores and the ferric reductase activity of roots, pH and acid phosphatases in the rhizosphere and bulk soil, and the secretion of acid phosphatases in roots. Our results revealed that shoot yields of peanut and maize plants were decreased by intercropping the plants, as compared to monocultured plants. Growing peanut plants in a mixture with maize, enhanced the shoot concentrations of Fe and Zn nearly 2.5-fold in peanut, while the Mn concentrations of peanut were little affected by intercropping. In the case of maize, the shoot concentrations of Fe, Zn and Mn were not significantly affected by intercropping with peanut. Intercropping also improved the shoot K concentration of peanut and maize, while it negatively affected the Ca concentration. In the intercropping of peanut/maize, the acid phosphatase activity of the rhizosphere and bulk soil and root secreted acid phosphatases were significantly higher than that of monocultured peanut and maize. In accordance, the shoot P concentrations of peanut and maize plants were much higher when they were intercropped with peanut or maize, respectively. The rhizosphere and bulk soil pH values were not clearly affected by different cropping systems. When compared to their monoculture treatments, the secretion of phytosiderophore from roots and the root ferric reducing capacity of the roots were either not affected or increased by 2-fold by the intercropping, respectively. The results indicate the importance of intercropping systems as a promising management practice to alleviate Fe deficiency stress. Intercropping also contributes to better nutrition of plants with Zn, P and K, most probably by affecting biological and chemical process in the rhizosphere.     

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.     

不同磷水平下玉米-大豆间作系统根系形态变化

本研究通过盆栽试验,探讨不同磷水平(0、50、100 mg P₂O₅·kg^-1,分别用P₀、P₅₀、P₁₀₀表示)下玉米与大豆间作系统根系形态的变化及其与磷吸收的关系,以明确玉米-大豆间作系统促进磷吸收的作用机制。结果表明: 不同磷水平下,间作显著改变了玉米和大豆的根系形态参数,提高了大豆根冠比。与单作模式相比,间作使玉米和大豆的根长、根表面积、根体积、根系干重分别显著增加25.6%、22.0%、39.2%、34.3%和28.1%、29.7%、37.3%、62.3%,而平均根直径分别显著降低15.2%和11.7%。不同磷水平下,磷素吸收当量比(LER_P)>1,玉米-大豆间作具有明显的磷吸收优势,且LER_P不受磷水平调控。间作诱导根系形态改变与磷吸收增加密切相关,其中玉米根系表面积增大、大豆根系长度增加是驱动玉米-大豆间作系统磷高效吸收的主要机制。根据回归方程,玉米根表面积和大豆根系长度增大10%,磷吸收量提高5%～10%。因此,与中等施磷水平(P₁₀₀)下的单作相比,玉米-大豆间作条件下磷肥减施1/2(P₅₀)并未降低玉米的磷吸收量。综上,玉米-大豆间作体系在减施磷肥条件下具有维持作物磷吸收的潜力。     

Maize root responses to drought stress depend on root class and axial position

In this study we tested the hypotheses that root classes would exhibit distinctive anatomical and architectural responses to drought stress, and that those responses would vary along the root axes. The root systems of four maize (Zea mays L.) sweet corn genotypes designated SC1, SC2, SC3 and SC4 were phenotyped under well-watered and drought treatments in greenhouse mesocosms, permitting increasing stratification of moisture availability as the drought progressed. Anatomical and architectural responses to drought were evaluated for each root class. Lignin distribution was assessed by image processing of UV-illuminated root cross-sections acquired by laser ablation tomography. The two cultivars with less biomass reduction under drought, SC3 and SC4, substantially enhanced lateral root development along the apical segments of axial roots when plants were grown with drought stress. These segments grew into the deeper part of the mesocosm where more moisture was available. Apical segments of the axial and large lateral roots from drought-stressed plants were thicker and had greater theoretical axial water conductance than basal segments, especially in SC3 and SC4. Basal segments of crown roots of SC3 and SC4 showed increased lignification of the stele under drought. Root anatomical and architectural responses to drought are complex and vary among cultivars and root classes, and along root axes. Drought-induced proliferation of lateral roots on apical segments of axial roots would be expected to enhance deep water acquisition, while lignification of axial roots could help preserve axial water transport.

     

Studies on the improvement in iron nutrition of peanut by intercropping with maize on a calcareous soil

Both rhizobox and field experiments were conducted to investigate nutritional interactions between peanut and maize in intercropping systems for Fe acquistion. Field observations indicated that Fe deficiency chlorosis symptoms in peanut grown in monoculture were more severe and widespread compared to those of peanuts intercropped with maize. This indicated a marked improvement in the iron nutrition of peanut intercropped with maize in the field and was further studied. In experiments with rhizoboxes, roots of maize and peanut were either allowed to interact with each other or prevented from making contact by inserting a solid plate between the root systems of the two species. A field experiment for four cropping treatments were examined: peanut grown separately in monoculture, normal peanut/maize intercropping, peanut/maize intercropping with solid plates between the root systems of the two crop species and peanut/maize intercropping with 30 μm nylon nets between the root systems. The results show that the chlorophyll and HCl-extractable Fe concentrations in young leaves of peanut in the intercropping system with unrestricted interactions of the roots of both plant species were much higher than those of peanut in monoculture. In the nylon mesh treatment, the beneficial effects of the maize extended to row 3. The improvement of Fe nutrition in the intercropping system got reduced but not diminished completely in the treatment with nylon net. It is suggested that the improvement in the Fe nutrition of peanut intercropped with maize was mainly caused by rhizosphere interactions between peanut and maize. This revised version was published online in June 2006 with corrections to the Cover Date.     

Reassessment of root competition for P of field-grown maize in pure and mixed cropping

Root competition for phosphate (P) in the field was calculated with maize i) as singly grown plants, ii) in pure cropping and iii) in mixed cropping with lupin. The experimentally determined parameters used for this purpose were thein situ root-distance pattern as registered by autoradiography and the P-depletion cylinder around a maize root as determined autoradiographically under radial diffusional flow. The results do not indicate any competition for P between roots of adjacently grown plants in either pure or mixed cropping. On the other hand, the P-depletion cylinders of approximately one third of the roots of an individual maize plant were found to overlap. However, when allowance was made for the concentration gradients within a depletion cylinder, the actual competition for phosphate was less than 1%.     

Root halotropism: Salinity effects on Bassia indica root

Abstract

Plant roots are responsible for the acquisition of nutrients and water from the soil and have an important role in plant response to soil stress conditions. The direction of root growth is gravitropic in general. Gravitropic responses have been widely studied; however, studies about other root tropisms are scarce. Soil salinity is a major environmental response factor for plants, sensed by the roots and affecting the whole plant. Our observations on root architecture of Kochia (Bassia indica) indicated that salinity may cue tropism of part of the roots toward increasing salt concentrations. We termed this phenomenon “positive halotropism”. It was observed that Kochia individuals in the field developed horizontal roots, originating from the main tap root, which was growing toward saline regions in the soil. Under controlled conditions in greenhouse experiments, Kochia plants were grown in pots with artificial soil salinity gradients, achieved by irrigation with saline and fresh water. It was shown that plants grown in low‐salt areas developed a major horizontal root toward the higher salt concentration region in the gradient. In regions of high salinity and in the absence of a salinity gradient, roots grew vertically without a major horizontal root. The novel finding of “positive halotropism” is discussed.     

Root and hyphal interactions influence N transfer by arbuscular mycorrhizal fungi in soybean/maize intercropping systems

In maize-soybean intercropping systems, the transfer of N from soybean to maize gives the intercropping system the advantage of improved N utilization and higher yields. Mycorrhiza acts as an important pathway for N transfer, providing a constant supply of N to sustain the growth and development of maize in its early stages. However, it is not clear how arbuscular mycorrhizal fungi (AMF) drive the transfer of N from soybean to maize in the intercropping system. Therefore, we quantified the amount of N transferred from soybean to maize under low and high N levels in the intercropping system, and the abundance and diversity of AMF involved in N transfer (¹⁵N-AMF) under different conditions by ¹⁵N leaf marker and DNA-SIP technology. We found that the interaction between roots and reducing the application of N fertilizer increased the amount of N transfer from soybean to maize. Compared with plastic plate separation (PS), no separation (NS) and mesh separation (MS) significantly increased the N fixation rate (from 14.33% to 39.09%), and the amount of N transfer under NS was 1.95–3.48 times that under MS. N transfer from soybean to maize ranged from 9.7 to 43.42 mg per pot in the no N treatment, while the addition of N fertilizer reduced N transfer by 14.12–66.28%. This is due to root interaction and reduced N fertilization increased the abundance and diversity of the ¹⁵N-AMF community, thereby promoting AMF colonization of maize and soybean roots. AMF colonization in soybean and maize roots under NS treatment was 6.47–17.24% higher than under MS treatment in all three levels of N addition. The increase of mycorrhiza in root system increased the N transfer from soybean to maize significantly. These results suggest that reduced N fertilizer in maize-soybean intercropping systems can increase N transfer by the mycorrhizal pathway, meeting maize N requirements and reducing chemical N fertilizer, which is important for sustainable agricultural development.     

不同种植模式对作物根系生长、产量及根际土壤微生物数量的影响

采用多年大田试验研究了小麦-大豆(A1)、小麦-甘薯(A2)、玉米(A3)、小麦/玉米/大豆(A4)和小麦/玉米/甘薯(A5)5种种植模式的根际环境变化特征和根系生长特性.结果表明:与A1、A2、A3和A5相比,A4提高了小麦、玉米、大豆在开花期和成熟期的生物量、根系活力和根干质量,提高了各作物根际土壤细菌、真菌和放线菌数量.各种植模式之间,植株生物量和根际微生物数量的变化规律为套作＞单作、大豆茬口＞甘薯茬口、边行＞中行.小麦/玉米/大豆(A4)套作模式通过改善3种作物的根际环境,促进了作物地下部根系生长和地上部生物量的增加,从而实现作物增产.     

Phosphorus sources and availability modify growth and distribution of root clusters and nodules of native Australian legumes

A variety of native Western Australian legumes produced root clusters in sand culture confirming field and published observations. In general, these legumes grew equally well when supplied with organic or inorganic sources of phosphorus. The nitrogen content of shoots and roots varied little among treatments for all species, however, phosphorus content was always greater in plants supplied with inositol‐P. The plasticity of root growth in response to localized placement of organic and inorganic sources of phosphorus was demonstrated using a simple ‘split root’ technique. Total root dry weight was, on average, more than doubled in P‐amended sand when compared with non‐amended sand. Root clusters tended to be produced in areas of relatively high phosphorus concentration and nodules in areas of low phosphorus concentration. Levels of phosphorus in lateral roots grown in P‐amended sand were significantly different from lateral roots grown in the corresponding non‐amended sand. Growth increases averaging 70% for white sand to over 100% for yellow sand indicated a large degree of ‘plasticity’ in roots under conditions of heterogeneous supply of phosphorus. Spatially exclusive development of organs for the acquisition of nutrients is discussed in relation to requirements for carbon in organ production and maintenance.     

Root distribution and interactions between intercropped species

Even though ecologists and agronomists have considered the spatial root distribution of plants to be important for interspecific interactions in natural and agricultural ecosystems, few experimental studies have quantified patterns of root distribution dynamics and their impacts on interspecific interactions. A field experiment was conducted to investigate the relationship between root distribution and interspecific interactions between intercropped plants. Roots were sampled twice by auger and twice by the monolith method in wheat (Triticum aestivum L.)/maize (Zea mays L.) and faba bean (Vicia faba L.)/maize intercropping and in sole wheat, maize, and faba bean up to 100 cm depth in the soil profile. The results showed that the roots of intercropped wheat spread under maize plants, and had much greater root length density (RLD) at all soil depths than sole wheat. The roots of maize intercropped with wheat were limited laterally, but had a greater RLD than sole-cropped maize. The RLD of maize intercropped with faba bean at different soil depths was influenced by intercropping to a smaller extent compared to maize intercropped with wheat. Faba bean had a relatively shallow root distribution, and the roots of intercropped maize spread underneath them. The results support the hypotheses that the overyielding of species showing benefit in the asymmetric interspecific facilitation results from greater lateral deployment of roots and increased RLD, and that compatibility of the spatial root distribution of intercropped species contributes to symmetric interspecific facilitation in the faba bean/maize intercropping. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Plant architectural responses in simultaneous maize/soybean strip intercropping do not lead to a yield advantage

Maize/soybean strip intercropping is a commonly used system throughout China with high crop yields at reduced nutrient input compared to sole maize. Maize is the taller crop, and due to its dominance in light capture over soybean in the intercrop, maize is expected to outperform maize in sole cropping. Conversely, soybean is the subordinate crop and intercropped soybean plants are expected to perform worse than sole soybean. Crop plants show plastic responses in plant architecture to their growing conditions to forage for light and avoid shading. There is little knowledge on plant architectural responses to growing conditions in simultaneous (non-relay) intercropping and their relationship to species yields. A two-year field experiment with two simultaneous maize/soybean intercropping systems with narrow and wide strips was conducted to characterise architectural traits of maize and soybean plants grown as intercrop and sole crops. Intercropped maize plants, especially those in border rows, had substantially greater leaf area, biomass and yield than maize plants in sole crops. Intercropped soybean plants, especially those in border rows, had lower leaf area, biomass and yield than sole soybean plants. Overall intercrop performance was similar to that of sole crops, with the land equivalent ratio (LER) being only slightly greater than one (1.03–1.08). Soybean displayed typical shade avoidance responses in the intercrop, such as greater internode elongation and changes in specific leaf area, but these responses could not overcome the consequences of the competition with the taller maize plants. Therefore, in contrast to relay intercrop systems, in the studied simultaneous maize/soybean system, plastic responses did not contribute to practically relevant increases in resource capture and yield at whole system (i.e., intercrop) level.