不同禾本科作物与花生混作对花生根系质外体铁的累积和还原力的影响

采用土培盆栽方法模拟玉米／花生、大麦／花生、燕麦／花生、小麦／花生、高粱／花生5种种植方式,研究混作对花生根系质外体铁的累积和还原力的影响．结果表明,当花生与5种分泌植物铁载体能力不同的禾本科作物混作时,花生新叶叶色正常,而单作花生则表现出严重的缺铁黄化症状,混作花生各部位的含铁量明显增加．与麦类作物(大麦、燕麦、小麦)混作的花生其各部位铁含量高于与玉米、高粱混作的花生,说明麦类作物改善花生铁营养的能力强于玉米、高粱,而两个玉米品种之间的能力差异不大。这主要是由于麦类作物分泌植物铁载体能力高于玉米、高粱．在花生生长至第50、60和70d时,混作花生根系质外体铁含量也随着逐渐增加,并始终高于单作花生．同时,混作明显地提高了花生根际土壤有效铁的含量,花生根系还原力也逐步提高．混作花生逐渐提高的还原力和介质中不断供给的易被花生还原吸收的铁。在改善花生的铁营养方面起了重要的作用．     

不同禾本科作物与花生混作对花生根系质外体铁的累积和还原力的影响

采用土培盆栽方法模拟玉米/花生、大麦/花生、燕麦/花生、小麦/花生、高粱/花生5种种植方式,研究混作对花生根系质外体铁的累积和还原力的影响.结果表明,当花生与5种分泌植物铁载体能力不同的禾本科作物混作时,花生新叶叶色正常,而单作花生则表现出严重的缺铁黄化症状,混作花生各部位的含铁量明显增加.与麦类作物(大麦、燕麦、小麦)混作的花生其各部位铁含量高于与玉米、高粱混作的花生,说明麦类作物改善花生铁营养的能力强于玉米、高粱,而两个玉米品种之间的能力差异不大,这主要是由于麦类作物分泌植物铁载体能力高于玉米、高粱.在花生生长至第50、60和70d时,混作花生根系质外体铁含量也随着逐渐增加,并始终高于单作花生.同时,混作明显地提高了花生根际土壤有效铁的含量,花生根系还原力也逐步提高.混作花生逐渐提高的还原力和介质中不断供给的易被花生还原吸收的铁,在改善花生的铁营养方面起了重要的作用.     

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.     

Molecular evidence for phytosiderophore‐induced improvement of iron nutrition of peanut intercropped with maize in calcareous soil

Peanut/maize intercropping is a sustainable and effective agroecosystem that evidently enhances the Fe nutrition of peanuts in calcareous soils. So far, the mechanism involved in this process has not been elucidated. In this study, we unravel the effects of phytosiderophores in improving Fe nutrition of intercropped peanuts in peanut/maize intercropping. The maize ys3 mutant, which cannot release phytosiderophores, did not improve Fe nutrition of peanut, whereas the maize ys1 mutant, which can release phytosiderophores, prevented Fe deficiency, indicating an important role of phytosiderophores in improving the Fe nutrition of intercropped peanut. Hydroponic experiments were performed to simplify the intercropping system, which revealed that phytosiderophores released by Fe‐deficient wheat promoted Fe acquisition in nearby peanuts and thus improved their Fe nutrition. Moreover, the phytosiderophore deoxymugineic acid (DMA) was detected in the roots of intercropped peanuts. The yellow stripe1‐like (YSL) family of genes, which are homologous to maize yellow stripe 1 (ZmYS1), were identified in peanut roots. Further characterization indicated that among five AhYSL genes, AhYSL1, which was localized in the epidermis of peanut roots, transported Fe(III)–DMA. These results imply that in alkaline soil, Fe(III)–DMA dissolved by maize might be absorbed directly by neighbouring peanuts in the peanut/maize intercropping system.     

玉米/花生混作改善花生铁营养对花生根瘤碳氮代谢及固氮的影响

研究了石灰性土壤上玉米 (Zea mays L.) /花生 (Arachishypogaea L.)混作改善花生铁营养对花生光合速率、光合产物的运输、花生各部位糖类含量、固氮酶活性以及根瘤内碳氮代谢及其有关酶活性的影响。结果表明 ,玉米 /花生混作改善花生铁营养能够明显增强固氮酶活性 ,进而增加了间作花生根瘤氨基酸的含量 ,这主要是由于玉米 /花生混作改善花生铁营养促进了花生光合作用 ,提高光合产物数量 ,增加光合产物由地上部向地下部的运输 ,但是处理间花生根瘤蔗糖和可溶性糖含量变化不大 ,单作花生根瘤还积累较多淀粉 ,说明不是光合产物的供应导致了花生固氮活性的差异。玉米 /花生混作对花生根瘤碳水化合物代谢水平影响较大 ,混作花生根瘤异柠檬酸脱氢酶 (IDH)、苹果酸脱氢酶 (MDH)、琥珀酸脱氢酶活性明显高于单作 ,而磷酸烯醇丙酮酸羧激 (PEPCK)活性低于单作花生 ,表明混作花生根瘤内三羧酸循环代谢水平较高 ,形成类菌体直接吸收利用的能量物质苹果酸和琥珀酸多 ,能够满足类菌体的固氮需求 ,因此 ,玉米 /花生混作改善花生铁营养增强根瘤碳水化合物代谢水平是提高花生固氮作用的重要原因之一     

玉米、小麦与花生间作改善花生铁营养机制的探讨

采用土培盆栽方法模拟研究了玉米／花生、小麦／花生间作对花生铁营养状况的影响及其作用机制。结果表明,禾本科作物与花生间作对花生的铁营养状况有显著影响：当花生与玉米或小麦分别间作时,花生新叶叶色正常,而花生单作则表现出严重的缺铁黄化现象,间作花生新叶活性铁、叶绿素含量明显高于单作,两种间作花生各部位铁含量和吸收量明显高于单作,间作明显地促进了铁向花生地上部的转移;在单作花生表现缺铁症状１４ｄ的时间范围内,其根系质外体铁含量仅是间作花生的５２％～８０％;而根系还原力则是单作花生在表现缺铁症状后迅速提高,至缺铁第６ｄ时还原力达到最大值,随后花生根系还原力迅速下降,而间作花生在０～１４ｄ内还原力增加速度缓慢,在１０～１４ｄ中其根系还原力明显地高于单作花生根系还原力。其主要原因可能是禾本科作物玉米、小麦根系分泌物（如：麦根酸类植物铁载体）螯合土壤中难溶性铁并被花生吸收利用。     

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.     

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.     

Interspecific root interactions and rhizosphere effects on salt ions and nutrient uptake between mixed grown peanut/maize and peanut/barley in original saline-sodic-boron toxic soil

Two glasshouse studies were conducted to investigate the effect of interspecific complementary and competitive root interactions and rhizosphere effects on the concentration and uptake of Na, Cl and B, and N, P, K, Ca, Mg, Fe, Zn and Mn nutrition of mixed cropped peanut with maize (Experiment I), and barley (Experiment II) grown in nutrient-poor saline-sodic and B toxic soil. Mixed cropped plants were grown in either higher density or lower density. The results of the experiment revealed that dry shoot weight decreased in peanut but increased in maize and barley with associated plant species compared to their monoculture. Shoot Na and Cl concentrations of peanut decreased significantly in both experiments, regardless of higher or lower density. The concentrations of Na also decreased in the shoots of mixed cropped maize and barley, but Cl concentrations increased slightly. The concentration of B significantly decreased in mixed cropping in all plant species regardless of higher or lower density. Rhizosphere chemistry was strongly and differentially modified by the roots of peanut, maize and barley, and mixed growing. There were significant correlations between the root-secreted acid phosphatases (S-APase), acid phosphatase in rhizosphere (RS-APase) and rhizosphere P concentration (RS-P) in the both experiments. The Fe-solubilizing activity (Fe-SA) and ferric reducing (FR) capacity of the roots were generally higher in mixed culture relative to their monoculture, which improved Fe, Zn and Mn nutrition of peanut. Further, there were also significant correlations among FR, Fe-SA and RS-Fe concentrations. Peanut facilitated P nutrition of maize and barley, while maize and barley improved K, Fe, Zn and Mn nutrition of peanut grown in nutrient-poor saline-sodic and B toxic soil.     

AhNRAMP1 iron transporter is involved in iron acquisition in peanut

Peanut/maize intercropping is a sustainable and effective agroecosystem to alleviate iron-deficiency chlorosis. Using suppression subtractive hybridization from the roots of intercropped and monocropped peanut which show different iron nutrition levels, a peanut gene, AhNRAMP1, which belongs to divalent metal transporters of the natural resistance-associated macrophage protein (NRAMP) gene family was isolated. Yeast complementation assays suggested that AhNRAMP1 encodes a functional iron transporter. Moreover, the mRNA level of AhNRAMP1 was obviously induced by iron deficiency in both roots and leaves. Transient expression, laser microdissection, and in situ hybridization analyses revealed that AhNRAMP1 was mainly localized on the plasma membrane of the epidermis of peanut roots. Induced expression of AhNRAMP1 in tobacco conferred enhanced tolerance to iron deprivation. These results suggest that the AhNRAMP1 is possibly involved in iron acquisition in peanut plants.     

与玉米混作改善花生铁营养对其根瘤形态结构及豆血红蛋白含量的影响

通过土培方法研究了与玉米混作对花生根瘤形态结构及固氮功能的影响。结果表明,玉米与花生混作能够明显地改善花生铁营养、提高根瘤豆血红蛋白的含量。同时,单作花生根瘤细胞液泡化程度较高,正在发育的根瘤细胞内类菌体数量明显地比混作的花生低。成熟根瘤细胞类菌体周膜外空间(细胞壁以内、周膜外的空间)体积变大。说明单作花生固氮酶活性较低的原因是缺铁抑制了豆血红蛋白的合成和改变了根瘤形态结构以及类菌体的超微结构。     

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.     

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.     

Root-microbial effects on plant iron uptake from siderophores and phytosiderophores

Collaborative experiments were conducted to determine whether microbial populations associated with plant roots may artifactually affect the rates of Fe uptake and translocation from microbial siderophores and phytosiderophores. Results showed nonaxenic maize to have 2 to 34-fold higher Fe-uptake rates than axenically grown plants when supplied with 1 μM Fe as either the microbial siderophore, ferrioxamine B (FOB), or the barley phytosiderophore, epi-hydroxymugineic acid (HMA). In experiments with nonsterile plants, inoculation of maize or oat seedlings with soil microorganisms and amendment of the hydroponic nutrient solutions with sucrose resulted in an 8-fold increase in FOB-mediated Fe-uptake rates by Fe-stressed maize and a 150-fold increase in FOB iron uptake rates by Fe-stressed oat, but had no effect on iron uptake by Fe-sufficient plants. Conversely, Fe-stressed maize and oat plants supplied with HMA showed decreased uptake and translocation in response to microbial inoculation and sucrose amendment. The ability of root-associated microorganisms to affect Fe-uptake rates from siderophores and phytosiderophores, even in short-term uptake experiments, indicates that microorganisms can be an unpredictable confounding factor in experiments examining mechanisms for utilization of microbial siderophores or phytosiderophores under nonsterile conditions.     

AhDMT1, a Fe<Superscript>2+</Superscript> transporter,is involved in improving iron nutrition and N<Subscript>2</Subscript> fixation in nodules of peanut intercropped with maize in calcareous soils

Peanut (Arachis hypogaea L.) is an important legume providing edible proteins and N₂ fixation. However, iron deficiency severely reduces peanut growth in calcareous soils. The maize/peanut intercropping effectively improves iron nutrition and N₂ fixation of peanut under pot and field conditions on calcareous soils. However, little was known of how intercropping regulates iron transporters in peanut. We identified AhDMT1 as a Fe²⁺ transporter which was highly expressed in mature nodules with stronger N₂ fixation capacity. Promoter expression analysis indicated that AhDMT1 was localized in the vascular tissues of both roots and nodules in peanut. Short-term Fe-deficiency temporarily induced an AhDmt1 expression in mature nodules in contrast to roots. However, analysis of the correlation between the complex regulation pattern of AhDmt1 expression and iron nutrition status indicated that sufficient iron supply for long term was a prerequisite for keeping AhDmt1 at a high expression level in both, peanut roots and mature nodules. The AhDmt1 expression in peanut intercropped with maize under 3 years greenhouse experiments was similar to that of peanut supplied with sufficient iron in laboratory experiments. Thus, the positive interspecific effect of intercropping may supply sufficient iron to enhance the expression of AhDmt1 in peanut roots and mature nodules to improve the iron nutrition and N₂ fixation in nodules. This study may also serve as a paradigm in which functionally important genes and their ecological significance in intercropping were characterized using a candidate gene approach.     

玉米/花生间作对土壤微生物和土壤养分状况的影响

通过大田试验,研究了玉米/花生间作对玉米和花生根区土壤微生物和土壤养分状况的影响.结果表明：与单作相比,间作能显著提高玉米和花生根区的土壤细菌数量;间作花生根区土壤真菌和放线菌数量与单作无显著差异;间作玉米根区土壤真菌和放线菌数量比单作明显提高;间作作物根区微生物群落功能多样性和代谢活性比单作有所改善.玉米/花生间作不同程度提高了整个间作系统根区的土壤碱解氮、速效磷、有机质含量及EC值,其中,间作玉米根区土壤养分的增加更为明显,说明玉米/花生间作可以较明显地改善两种作物根区的微生物和养分状况,土壤微生态环境的改善又会促进作物地上部的生长.     

花生与药材套种对土壤微生物区系的影响

通过茅苍术、京大戟、黄姜（盾叶薯蓣）、半夏和阔叶麦冬5种药用植物与花生套种的盆栽试验,测定了不同时期花生土壤中的细菌、放线菌、霉菌和酵母菌数量,探讨利用药用植物套种花生缓解花生连作障碍的可行性.结果表明：茅苍术组和京大戟组抑制土壤霉菌效果最好,在花生花针期分别比对照组减少了53.87%和29.59%;在花生收获后的土壤中霉菌数量增加,有利于物质循环和养分还田.茅苍术、京大戟和半夏组土壤中细菌数量增加. 在花生花针期,5种药用植物套作组土壤中酵母菌数量均高于对照.霉菌形态鉴定结果表明,在茅苍术、京大戟和黄姜套作组中均未检测出花生常见病原菌.药用植物和花生套种可以有效调节土壤微生物区系.     

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.     

间套作体系作物-土壤铁和锌营养研究进展

间套作能够提高对土地、光、水和养分等资源的利用,对增加农田生物多样性、发展可持续生态农业具有重要的理论和实践意义.间套作有助于改善土壤中微量元素铁和锌的活化、吸收和植株体内微量元素营养状况,实现微量元素的生物强化.本文综述了禾本科/双子叶植物间套作（主要以玉米/花生间作为例）对双子叶植物铁和锌营养的改善效应,从种间根际分子和生理过程、根系形态、分布及根际微生物变化方面详细阐述了可能的机理,同时综述了铁和锌在间套作植株体内累积、转运和分配方面的研究进展,对影响间套作体系铁和锌营养的可能因素进行了简单分析.从土壤养分活化吸收、植株体内养分累积分配和栽培管理措施及环境因子等方面指出目前研究中存在的问题,提出以后应加强间套作体系铁和锌营养田间试验、定量化、动态性和系统化方面的研究.