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。     

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。     

小麦-蚕豆间作对根系分泌低分子量有机酸的影响

通过盆栽试验收集了不同生育期单作和间作小麦、蚕豆的根系分泌物,用HPLC分析了根系分泌物中低分子量有机酸的含量和种类.结果表明: 小麦-蚕豆间作显著提高了有机酸的分泌量,在小麦分蘖期(57 d)、孕穗期(120 d)和灌浆期(142 d),间作使小麦根系有机酸分泌量分别提高155%、35.6%和92.6%;在蚕豆分枝期(57 d)和籽粒膨大期(142 d),间作使蚕豆根系有机酸分泌量提高87.4%和38.7%.小麦-蚕豆间作改变了根系分泌物中有机酸的种类,与单作小麦相比,在分蘖期,间作小麦根系分泌物中增加了乳酸;在拔节期(98 d),间作小麦根系分泌物中增加了柠檬酸,但未检测到乙酸;在蚕豆分枝期,间作蚕豆根系分泌物中增加了乙酸,但未检测到乳酸;在蚕豆籽粒膨大期,间作蚕豆根系分泌物中增加了乳酸.小麦-蚕豆间作提高了小麦根系有机酸的分泌速率,在小麦孕穗期,间作小麦分泌柠檬酸、富马酸的速率是单作小麦的179和184倍;在小麦灌浆期,间作小麦分泌乳酸的速率是单作的2.53倍.总之,小麦-蚕豆间作增加了有机酸的分泌量,改变了根系分泌物中有机酸的种类,提高了小麦根系有机酸的分泌速率.     

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。     

施氮对间作蚕豆根际微生物区系和枯萎病发生的影响

通过田间小区试验,研究了小麦、蚕豆间作条件下4个施氮水平(0,56.25,112.5kg.hm-2和168.75kg.hm-2)对蚕豆根际微生物区系和蚕豆枯萎病发生的影响。结果表明,单作和间作条件下,施氮显著增加了蚕豆根际的微生物数量,在N2(112.5kg.hm-2)水平下达到最高值;施氮对土壤微生物多样性无显著影响,但减轻了单、间作蚕豆枯萎病的发生,且在N2水平下发病最轻。与单作相比,间作显著增加了蚕豆根际的细菌、真菌、放线菌数量、微生物总数和微生物多样性,尤其在N0、N1(56.25kg.hm-2)和N2(112.5kg.hm-2)水平下间作对蚕豆根际微生物的促进效应明显,且以真菌和放线菌的增幅较大.N0、N1和N2水平下间作显著降低了蚕豆枯萎病的发病率和病情指数。小麦蚕豆间作下适量施氮能有效调节蚕豆根际微生物区系,是抑制蚕豆枯萎病发生的有效措施。     

核桃-小麦复合系统中细根生长动态及竞争策略

以核桃(Juglans regia)-小麦(Triticum aestivum)间作复合系统为研究对象,用微根窗和根钻相结合的方法采样,研究复合系统中核桃和小麦细根年内年际的生长动态和竞争适应策略,为农林复合系统的经营管理和竞争模型的建立提供理论依据和技术支持。结果表明,间作核桃和小麦根系均在上半年有一个大的生长高峰(5月和4月),在下半年有一个小的生长高峰(9月和11月),二者的竞争主要发生在上半年的大生长高峰期。在各年份各土层,间作核桃的根长密度均低于单作核桃,且在从第7年开始存在显著差异。在0—20 cm土层间作小麦根长密度在第3—7年间获得迅速提高,从第7年开始显著高于单作小麦,但在20 cm以下土层则相反。间作使核桃和小麦细根生态位实现了分离,11年的观察期内间作核桃比单作核桃细根的垂直分布中心下移了6.59 cm,间作小麦比单作小麦的上移了8.59 cm。在根系竞争策略方面,小麦根系是通过短期内的快速生长,迅速占据土壤空间获得竞争优势;而核桃根系是通过根系的逐年积累,逐步占据土壤空间从而获得竞争优势。可以干扰核桃根系积累过程的"竞争-干扰-再平衡"农林复合经营管理策略可以让复合系统中核桃和小麦保持各自竞争优势的情况下实现共存。在根系形态方面,自身细根直径较小者小麦在剧烈竞争区域以增加细根直径减小比根长来适应竞争,而自身细根直径较大者核桃则相反。     

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.     

低磷胁迫下磷高效基因型大麦的根系形态特征

在根袋土培盆栽条件下,以磷高效基因型DH110+、DH147和低效基因型DH49大麦为试验材料,利用根系分析系统分析不同施磷(P₂O₅)水平(极低磷25 mg·kg^-1、低磷50 mg·kg^-1和正常磷75 mg·kg^-1)下,磷高效基因型大麦的根系形态特征及其与植株磷素吸收的关系.结果表明： 低磷胁迫显著降低大麦生物量和磷吸收量,其中磷高效基因型的生物量和磷吸收量在各施磷水平下分别为低效基因型的1.24～1.70和1.18～1.83倍;大麦的总根长、总根表面积、平均根系直径、不定根长及其根表面积、侧根长及其根表面积均随施磷水平的降低而显著降低,其中磷高效基因型大麦在各施磷水平下的总根长、总根表面积、比根长、侧根长及根表面积分别为低效基因型的1.46～2.06、1.12～1.51、1.35～1.72、1.69～2.42和1.40～1.78倍,而平均根系直径为低效基因型的70.6%～90.2%;主成分分析表明,平均根系直径、比根表面积和比根长受基因型差异的影响较为明显,是区分两类磷效率基因型大麦根系形态差异的主要指标;偏最小二乘回归分析表明,各施磷水平下,总根长、总根表面积对大麦植株磷素吸收贡献均较大,随施磷水平降低,不定根长、不定根表面积对大麦植株磷素吸收的贡献明显降低,而平均根系直径、比根长、侧根长及其根表面积的贡献明显增加.磷高效基因型大麦可通过维持侧根的生长、根细度和比根长的增加来适应低磷胁迫.     

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.     

磷高效利用杉木对低磷胁迫的适应性与内源激素的相关性

激素是植物适应逆境的重要信号物质, 从激素调控角度研究植物对养分匮乏环境的适应机制对磷高效营养基因型的选育具有重要意义。该研究通过分析被动忍受型(M1)与主动活化型(M4)两个磷高效利用杉木(Cunninghamia lanceolata)基因型在低磷胁迫下不同处理阶段的激素含量变化规律, 结合根系形态变化、干物质及养分分配规律, 研究磷高效利用杉木对低磷胁迫的适应性与内源激素的相关性。结果表明: 低磷处理下, 磷高效利用杉木M1与M4叶的激素含量与其适应特性之间无相关性, 而根系的激素含量与根系生长显著相关。低磷处理条件下, M1与M4根系中的IAA含量自27 h起表现为大于高磷对照, 且随时间延长呈增加趋势。根系中的IAA含量与根表面积、体积及根长等显著正相关, IAA的增加诱导了根系的增长, M1与M4均表现出一定的根系增长量。其中, M4存在明显的IAA由地上向基部积累的现象, M4的根系增生能力比M1更强。同时, 根系增长促使更多的干物质分配到根系, M4的根冠比在整个处理过程中均高于高磷对照。与IAA相同, M1与M4根系的ABA与GA₃含量总体也表现为低磷处理>高磷对照, 但随时间延长, 低磷条件下ABA与GA₃的含量呈下降趋势, 二者与根系增长量呈负相关关系。M1与M4根系内的ZT含量在低磷条件下也呈下降趋势, 且逐渐低于高磷对照, 而其与低磷适应特性间并无显著相关性。可见, 低磷胁迫下, 磷高效利用杉木M1与M4根系中的IAA、ABA与GA₃含量与其根系形态变化密切相关, 各器官的物质、能量、信息的综合调控是植物适应低磷逆境的重要生存策略。     

Intercropping influences component and content change of flavonoids in root exudates and nodulation of Faba bean

Flavonoids produced by legume roots are signal molecules acting as nod gene inducers for the symbiotic rhizobium partner. Nevertheless, the changes of flavonoids in root exudates in intercropping system are still unknown. Based on pot experiment of faba bean and wheat intercropping, here we showed that faba bean and wheat intercropping increased the nodules number and dry weight, dry weight per nodule of faba bean compared with those found in monocropping, and the increase of faba bean nodulation was likely caused by the enhancement with flavonol, isoflavone, chalcone and hesperetin in its root exudates. It also promoted exudation of five types of flavonoids by wheat compared with monocropping. Our findings suggest that the flavonoids in root exudates have a positive effect on the nodulation and nitrogen fixation of faba bean in faba bean and wheat intercropping.     

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。     

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.     

基于芽苗砧嫁接油茶砧穗创伤后内源激素动态变化分析

为研究油茶(Camellia oleifera A)嫁接时穗条和砧木创伤后内源激素动态变化规律,解析影响砧穗嫁接面愈合的生理机制,为油茶砧穗愈合生长机理提供理论支持。以树龄6年的长林18号和53号的穗条和实生砧木为材料,按照芽苗砧嫁接方法切割穗条S0(0 min)、S10(10 min)、S40(40 min)和砧木茎段Z0(0 min)、Z10(10 min)后,利用液质联用法(HPLC-MS)测定吲哚乙酸(IAA)、脱落酸(ABA)、反式玉米素(TZR)、玉米素(Zeatin)、水杨酸(SA)和茉莉酸(JA)含量,分析不同时间段内源激素变化及品种间差异的关系。结果显示:创伤后18号的TZR、Zeatin和SA含量总体高于53号;18号IAA、SA和JA逐渐下降;TZR和Zeatin分别在S10和S0达最高值后下降;ABA在S10达最高值。53号IAA和JA爱S10达最高值后下降;TZR、Zeatin和SA在S10达低值后逐渐上升;ABA在S0达高值后逐渐下降。砧木茎段创伤前后激素含量除JA外18号高于53号;两品种Z0时激素含量下降,Z10后上升,仅53号ABA和SA含量在Z0达高值后下降。砧木茎段和根部激素含量在品种间除JA外18号高于53号,茎段的IAA、ABA高于根部,其他激素为根部高于茎段。激素比值在品种间和部位间差异明显;IAA/ABA、IAA/TZR、IAA/Zeatin和IAA/JA、ABA/TZR、ABA/Zeatin和ABA/JA比值为53号高于18号;穗条内SA/IAA为18号高于53号,SA/JA和SA/ABA为53号高于18号;砧木茎段均为18号高于53号;TZR/SA、TZR/JA比值在穗条和砧木茎段为18号高于53号。两品种创伤后IAA与JA极显著正相关,而IAA与SA,SA与JA在18号极显著正相关,53号极显著或显著负相关;53号TZR、Zeatin、SA间极显著或显著正相关,JA与TZR、Zeatin和SA极显著负相关。砧木茎段创伤后18号激素间为极显著或显著正相关;53号TZR和Zeatin与IAA、JA极显著正相关,与SA存在显著负相关,SA与JA有显著负相关。砧木茎段和根部间品种间仅在SA与各激素间相关性存在差异,其他激素间存在极显著正相关或负相关。综上所述,砧穗创伤后激素水平上18号在创伤面易于愈伤组织发育,而53号抗逆激素水平较高且与细胞分裂增殖类激素负相关,可能影响53号嫁接后愈合生长;嫁接应在创伤后10 min内较为适宜;砧穗间激素含量及比值的差异可能会影响后期嫁接部位形态重建以及穗条生长。     

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.     

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.