Inoculation with Azospirillum argentinense Az19 improves the yield of maize subjected to water deficit at key stages of plant development

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(a) Biofilm production of B cereus group strains in different materials and (b) from source isolation: light blue circles, powered infant formula; gray squares, surfaces; orange triangles, infant food; purple triangles, eggshell. *p≤0.05, ****p≤0.0001. In the X axis are the materials and, in the Y, the specific biofilm formation (SBF). Top Kruskal–Wallis post hoc Dunn's. Lower ANOVA." ] ] ] "autores" => array:1 [ 0 => array:2 [ "autoresLista" => "Itzel-Maralhi Cruz-Facundo, Roberto Adame-Gómez, Natividad Castro-Alarcón, Jeiry Toribio-Jiménez, Yaneth Castro-Coronel, María-Cristina Santiago-Dionisio, Marco-Antonio Leyva-Vázquez, David Tafolla-Venegas, Arturo Ramírez-Peralta" "autores" => array:9 [ 0 => array:2 [ "nombre" => "Itzel-Maralhi" "apellidos" => "Cruz-Facundo" ] 1 => array:2 [ "nombre" => "Roberto" "apellidos" => "Adame-Gómez" ] 2 => array:2 [ "nombre" => "Natividad" "apellidos" => "Castro-Alarcón" ] 3 => array:2 [ "nombre" => "Jeiry" "apellidos" => "Toribio-Jiménez" ] 4 => array:2 [ "nombre" => "Yaneth" "apellidos" => "Castro-Coronel" ] 5 => array:2 [ "nombre" => "María-Cristina" "apellidos" => "Santiago-Dionisio" ] 6 => array:2 [ "nombre" => "Marco-Antonio" "apellidos" => "Leyva-Vázquez" ] 7 => array:2 [ "nombre" => "David" "apellidos" => "Tafolla-Venegas" ] 8 => array:2 [ "nombre" => "Arturo" "apellidos" => "Ramírez-Peralta" ] ] ] ] "resumen" => array:1 [ 0 => array:3 [ "titulo" => "Highlights" "clase" => "author-highlights" "resumen" => "<ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005">•We observe in the strains from foods a higher production of biofilms in PVC.</li><li class="elsevierStyleListItem" id="lsti0010">•In the BHI broth, no submerged biofilms were found.</li><li class="elsevierStyleListItem" id="lsti0015">•The enterotoxin profile most common includes genes for the three enterotoxins.</li></ul>" ] ] ] "idiomaDefecto" => "en" "EPUB" => "https://multimedia.elsevier.es/PublicationsMultimediaV1/item/epub/S0325754123000226?idApp=UINPBA00004N" "url" => "/03257541/0000005500000003/v1_202309280755/S0325754123000226/v1_202309280755/en/main.assets" ] "itemAnterior" => array:18 [ "pii" => "S032575412300024X" "issn" => "03257541" "doi" => "10.1016/j.ram.2023.02.002" "estado" => "S300" "fechaPublicacion" => "2023-07-01" "aid" => "538" "copyright" => "Asociación Argentina de Microbiología" "documento" => "article" "crossmark" => 1 "subdocumento" => "fla" "cita" => "Rev Argent Microbiol. 2023;55:251-4" "abierto" => array:3 [ "ES" => false "ES2" => false "LATM" => false ] "gratuito" => false "lecturas" => array:1 [ "total" => 0 ] "en" => array:14 [ "idiomaDefecto" => true "cabecera" => "Brief report" "titulo" => "First nosocomial outbreak of SME-4-producing Serratia marcescens in South America" "tienePdf" => "en" "tieneTextoCompleto" => "en" "tieneResumen" => array:3 [ 0 => "en" 1 => "en" 2 => "es" ] "paginas" => array:1 [ 0 => array:2 [ "paginaInicial" => "251" "paginaFinal" => "254" ] ] "titulosAlternativos" => array:1 [ "es" => array:1 [ "titulo" => "Primer brote nosocomial por Serratia marcescens productora de SME-4 en Sudamérica" ] ] "contieneResumen" => array:2 [ "en" => true "es" => true ] "contieneTextoCompleto" => array:1 [ "en" => true ] "contienePdf" => array:1 [ "en" => true ] "resumenGrafico" => array:2 [ "original" => 0 "multimedia" => array:7 [ "identificador" => "fig0005" "etiqueta" => "Figure 1" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr1.jpeg" "Alto" => 982 "Ancho" => 1074 "Tamanyo" => 74318 ] ] "descripcion" => array:1 [ "en" => "Phenotypic detection of SME-4. Double disc-synergy test: boronic acid (300μg)–imipenem (10μg) and imipenem (30μg)–amoxicillin clavulanic acid (30μg)." ] ] ] "autores" => array:1 [ 0 => array:2 [ "autoresLista" => "Juana Vega, Carlos H. Rodriguez, Ignacio Viscardi, Carlos Vay, Silvina Torres, Emilce Tabares, Angela Famiglietti, Marcela Nastro" "autores" => array:8 [ 0 => array:2 [ "nombre" => "Juana" "apellidos" => "Vega" ] 1 => array:2 [ "nombre" => "Carlos H." "apellidos" => "Rodriguez" ] 2 => array:2 [ "nombre" => "Ignacio" "apellidos" => "Viscardi" ] 3 => array:2 [ "nombre" => "Carlos" "apellidos" => "Vay" ] 4 => array:2 [ "nombre" => "Silvina" "apellidos" => "Torres" ] 5 => array:2 [ "nombre" => "Emilce" "apellidos" => "Tabares" ] 6 => array:2 [ "nombre" => "Angela" "apellidos" => "Famiglietti" ] 7 => array:2 [ "nombre" => "Marcela" "apellidos" => "Nastro" ] ] ] ] "resumen" => array:1 [ 0 => array:3 [ "titulo" => "Highlights" "clase" => "author-highlights" "resumen" => "<ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005">•Serratia marcescens represents a nosocomial pathogen worldwide.</li><li class="elsevierStyleListItem" id="lsti0010">•First description of a nosocomial outbreak by SME-4 in South America.</li><li class="elsevierStyleListItem" id="lsti0015">•Class A carbapenemases different from KPC may be underestimated.</li></ul>" ] ] ] "idiomaDefecto" => "en" "EPUB" => "https://multimedia.elsevier.es/PublicationsMultimediaV1/item/epub/S032575412300024X?idApp=UINPBA00004N" "url" => "/03257541/0000005500000003/v1_202309280755/S032575412300024X/v1_202309280755/en/main.assets" ] "en" => array:21 [ "idiomaDefecto" => true "cabecera" => "Original article" "titulo" => "Inoculation with Azospirillum argentinense Az19 improves the yield of maize subjected to water deficit at key stages of plant development" "tieneTextoCompleto" => true "paginas" => array:1 [ 0 => array:2 [ "paginaInicial" => "255" "paginaFinal" => "261" ] ] "autores" => array:1 [ 0 => array:4 [ "autoresLista" => "Julia Elena García, Mónica Ruiz, Guillermo Andrés Maroniche, Cecilia Creus, Mariana Puente, Myriam Sara Zawoznik, María Daniela Groppa" "autores" => array:7 [ 0 => array:3 [ "nombre" => "Julia Elena" "apellidos" => "García" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "a" "identificador" => "aff0005" ] ] ] 1 => array:3 [ "nombre" => "Mónica" "apellidos" => "Ruiz" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "b" "identificador" => "aff0010" ] ] ] 2 => array:3 [ "nombre" => "Guillermo Andrés" "apellidos" => "Maroniche" "referencia" => array:2 [ 0 => array:2 [ "etiqueta" => "c" "identificador" => "aff0015" ] 1 => array:2 [ "etiqueta" => "d" "identificador" => "aff0020" ] ] ] 3 => array:3 [ "nombre" => "Cecilia" "apellidos" => "Creus" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "c" "identificador" => "aff0015" ] ] ] 4 => array:3 [ "nombre" => "Mariana" "apellidos" => "Puente" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "a" "identificador" => "aff0005" ] ] ] 5 => array:4 [ "nombre" => "Myriam Sara" "apellidos" => "Zawoznik" "email" => array:1 [ 0 => "myriamz@ffyb.uba.ar" ] "referencia" => array:2 [ 0 => array:2 [ "etiqueta" => "e" "identificador" => "aff0025" ] 1 => array:2 [ "etiqueta" => "*" "identificador" => "cor0005" ] ] ] 6 => array:3 [ "nombre" => "María Daniela" "apellidos" => "Groppa" "referencia" => array:1 [ 0 => array:2 [ "etiqueta" => "f" "identificador" => "aff0030" ] ] ] ] "afiliaciones" => array:6 [ 0 => array:3 [ "entidad" => "Instituto Nacional de Tecnología Agropecuaria (INTA), Instituto de Microbiología y Zoología Agrícola, Argentina" "etiqueta" => "a" "identificador" => "aff0005" ] 1 => array:3 [ "entidad" => "Instituto Nacional de Tecnología Agropecuaria (INTA), Estación Experimental San Juan, Argentina" "etiqueta" => "b" "identificador" => "aff0010" ] 2 => array:3 [ "entidad" => "Facultad de Ciencias Agrarias, Universidad Nacional de Mar del Plata, Argentina" "etiqueta" => "c" "identificador" => "aff0015" ] 3 => array:3 [ "entidad" => "Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina" "etiqueta" => "d" "identificador" => "aff0020" ] 4 => array:3 [ "entidad" => "Universidad de Buenos Aires, Facultad de Farmacia y Bioquímica, Buenos Aires, Argentina" "etiqueta" => "e" "identificador" => "aff0025" ] 5 => array:3 [ "entidad" => "Universidad de Buenos Aires, Instituto de Química y Fisicoquímica Biológicas (IQUIFIB), Facultad de Farmacia y Bioquímica, Buenos Aires, Argentina" "etiqueta" => "f" "identificador" => "aff0030" ] ] "correspondencia" => array:1 [ 0 => array:3 [ "identificador" => "cor0005" "etiqueta" => "⁎" "correspondencia" => "Corresponding author." ] ] ] ] "titulosAlternativos" => array:1 [ "es" => array:1 [ "titulo" => "La inoculación con Azospirillum argentinense Az19 mejora el rendimiento del maíz expuesto a déficit hídrico durante estadios clave del desarrollo vegetal" ] ] "resumenGrafico" => array:2 [ "original" => 0 "multimedia" => array:7 [ "identificador" => "fig0010" "etiqueta" => "Figure 2" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr2.jpeg" "Alto" => 959 "Ancho" => 3175 "Tamanyo" => 268312 ] ] "descripcion" => array:1 [ "en" => "Changes in maize height (A), percentage of empty ears (B), and yield (C) according to irrigation and inoculation treatments (field assay 2). Mean values and standard deviation determined at physiological maturity are shown. Sampling was performed as described in “Materials and methods” section. C: full-irrigated control; WR-Fl: water restriction at flowering; WR-Gf: water restriction at grain filling; GS: drought-sensitive genotype; GT: drought-tolerant genotype. Data was square root-transformed (B) or not transformed (A and C) and analyzed by two-way ANOVA plus the Tukey's test. Different letters indicate significant differences between treatments irrespective of the plant genotype (p˂0.1)." ] ] ] "textoCompleto" => "IntroductionIn Argentina, maize (Zea mays L.) was traditionally cultivated in the Undulating Pampa region (Pampa Ondulada) (32° to 35.8° S and 58° to 62° W), a temperate-humid region without climatic constraints and adequate for extensive agriculture. However, due to the massive cultivation of soybean in the last decades, maize production has been expanded outside that optimal zone to semiarid environments (<700mm precipitation per year), where high temperatures and water restrictions are relatively frequent events<a class="elsevierStyleCrossRef" href="#bib0260">20</a>.Flowering is a key phenological stage for cereal production, and water deficit occurring during that stage adversely impacts yield due to its negative effect on grain development. Water limitation at flowering leads to impaired reproductive performance, resulting in a decreased number of grains per hectare and decreased final grain weight<a class="elsevierStyleCrossRef" href="#bib0255">19</a>.In addition to plant breeding techniques, microbial biotechnology is increasingly positioned as a valuable strategy to improve crop performance under adverse conditions. In this context, the introduction of plant growth-promoting rhizobacteria (PGPR) is an environmental-friendly alternative to chemical treatments for sustainable production systems. PGPR comprise those microorganisms that colonize the rhizosphere of plants and promote plant growth. Within this group, the genus Azospirillum has been one of the most studied and is usually regarded as a model bacterium to study plant-bacteria interactions<a class="elsevierStyleCrossRef" href="#bib0295">27</a>. Azospirillum can interact with a wide range of plant species due to its recognized metabolic versatility<a class="elsevierStyleCrossRef" href="#bib0295">27</a>. For example, the strain Az39 of Azospirillum argentinense, used in most commercial formulations in Argentina, was tested in various cereals and was found to increase yield significantly<a class="elsevierStyleCrossRefs" href="#bib0195">7,9,30</a>.It has also been documented that plants inoculated with PGPR exhibit greater tolerance to abiotic stresses such as hydric and osmotic stresses<a class="elsevierStyleCrossRef" href="#bib0250">18</a>. The beneficial effects of PGPR inoculation in plants subjected to osmotic stress include biomass increase and water status improvement<a class="elsevierStyleCrossRef" href="#bib0245">17</a>. Under field conditions, wheat inoculation with A. baldaniorum Sp245 increased cell wall elasticity in the flag leaf and reduced yield losses due to moderate drought at the anthesis stage. In addition, this strain mitigated the negative effect of drought and led to increased grain yield and mineral content (Mg, K, and Ca) compared to non-inoculated plants<a class="elsevierStyleCrossRef" href="#bib0190">6</a>.Previously, we characterized the strain Az19 of A. argentinense from the PGPR collection of IMyZA INTA and observed that this strain had similar PGPR features but better attributes to cope with osmotic stress than A. argentinense Az39, the strain typically included in commercial inoculants<a class="elsevierStyleCrossRef" href="#bib0215">11</a>. Az19 is a collection strain isolated in 1982 from wheat field fallow. Since this strain has already demonstrated higher in vitro survival under saline and osmotic stress than Az39 and higher trehalose production<a class="elsevierStyleCrossRef" href="#bib0215">11</a>, we hypothesized that this bacterium could also display higher survival in the soil/root interface and thus contribute to sustain plant growth and yield under water deficit, which usually entails osmotic imbalances. The purpose of this study was to assess if seed inoculation with the strain Az19 ameliorates the negative effects of water restriction at specific phenological stages on the yield of a commercial maize hybrid to a greater extent than the Az39 strain (greenhouse experiment and field assay 1) and on different contrasting maize genotypes for their sensitivity to drought (field assay 2).Materials and methodsPlant materialTwo commercial maize hybrids, DOW 510 PW (greenhouse experiment) and DOW DS 515 PW (field experiment 1), and two maize genotypes obtained at INTA Pergamino, Argentina, LP 29×LP 2542 (drought-sensitive) and LP 882 (923)×LP 4703 (drought-tolerant) (field experiment 2), were used.Azospirillum argentinense strains used for maize inoculationAz39 and Az19 were obtained from the PGPR collection of IMyZA INTA Castelar. Az39 was used as the reference strain.Both A. brasilense strains were cultured in OAB liquid medium<a class="elsevierStyleCrossRef" href="#bib0285">25</a> supplemented with 1g/l NH4Cl (for an optimal bacterial growth) at 30°C in a rotary shaker at 180rpm for 48h. Viable cell number (CFU/ml) was quantified using the drop plate method<a class="elsevierStyleCrossRef" href="#bib0225">13</a> on plates containing Congo Red medium<a class="elsevierStyleCrossRef" href="#bib0300">28</a>.Greenhouse experimentSurface disinfected (ethanol 80% and sodium hypochlorite 4%, v/v) seeds of the maize hybrid DOW 510 PW were pregerminated in a wet chamber at 25°C, and uniform-developed seedlings were inoculated 24h later by immersion for 2h in an inoculum containing 5×107 colony-forming units (CFU) of the corresponding A. argentinense strain per seedling. A set of non-inoculated seedlings was included. Emerging seedlings were subsequently transferred to 10l plastic pots containing a mixture of sterile sand, soil, vermiculite, and perlite (3:3:3:1) as substrate.All treatments (including controls) were irrigated with Hoagland nutrient solution. Water restriction was applied once at V2 or twice at V2 and flowering, and plants were harvested at V3 or physiological maturity, respectively. Water deficit at V2 was achieved by withholding the irrigation for 10 days while water deficit at flowering was achieved by withholding irrigation for 20 days after flowering. Male and female flowering dates were recorded when 50% of the plants were shedding pollen or showed 1-cm long stigmas, respectively.A randomized complete block design consisting of 3 blocks with 12 plants per treatment was used. Length, fresh weight, and dry weight of the aerial part and the relative water content (RWC) of the last expanded leaf were measured at V3 (plants exposed to water deficit at V2). RWC was determined according to the method described by Barrs and Weatherley<a class="elsevierStyleCrossRef" href="#bib0170">2</a> based on the following formula:<elsevierMultimedia ident="eq0005"></elsevierMultimedia>At physiological maturity, the percentage of fertile plants (grain-producing plants), the ear weight, and the grain weight per plant were determined (plants exposed to water deficit at V2 and flowering).Field assaysBoth field assays were conducted in plots belonging to the agricultural experimental station of the Instituto Nacional de Tecnología Agropecuaria in San Juan province (INTA EEA San Juan), located near Pocito city (31°39′0″S, 68°33′0″W), during the maize campaign 2015/16. We selected this location because of the recognized scarcity of rainfall, which is appropriate to mimic a water-deficient condition requiring controlled irrigation.The soil of this area is classified as Entisol, and since 2009, these plots have been under maize production, with fallow between cycles. Plots received 200kg N/ha (100kg at sowing and 100kg at V7). The mean temperature at silking (±15 days) was 30.2°C, and from that stage to grain filling was 24.1°C. Accumulated rainfall in the same periods was 13.5 and 7.2mm, respectively.Field assay 1. The commercial maize hybrid DOW DS 515 PW was used. Before planting, seeds were inoculated with the corresponding A. argentinense strain (12ml/kg seeds; 1×109CFU/ml of inoculant) and then sown at a plant density of 70000plants/ha, with a distance of 0.63m between rows and 0.19m between plants. A split-plot experimental design was used, the main plots corresponding to factor A, water restriction, and the secondary plots to factor B, inoculation, with 5 repetitions (plots) per treatment. Each plot included eight 5-m long rows.Water restriction (factor A) included a full-irrigated control, irrigated at 100% of the calculated evapotranspiration (ETc) throughout the entire crop cycle, and two alternative water deficit treatments: one imposed at flowering, with the irrigation reduced to 25% of the ETc between −15 to +15 days respect to the flowering date (WR-Fl), and the other one imposed at grain filling, with the irrigation reduced to 25% of the ETc from grain set up to physiological maturity (WR-Gf). Inoculation treatments (factor B) were the same as in the greenhouse experiment: Az39, Az19, none (control).Male and female flowering dates were recorded when more than 15 plants per row were shedding pollen and had 1-cm long stigmas, respectively. Irrigation was performed via a surface drip irrigation system with hoses of 1.27cm in diameter and emitters every 25cm, with a flow of 2.5l/h. The irrigation rate was established based on the calculated crop evapotranspiration. Yield (kg/ha) and empty ear rates (%) were evaluated at physiological maturity by sampling two random rows. The plant stand was uniform across treatments.Field assay 2. Only strain Az19 was used as inoculation treatment, and two non-commercial hybrids provided by the Breeding Program of INTA EEA Pergamino, Argentina, were tested in this trial: the drought-sensitive genotype LP 29×LP 2542 and the drought-tolerant genotype LP 882 (923)×LP 4703. Water restriction was imposed as in field assay 1, and yield and empty ear rate were evaluated as described. Plant height was an additional parameter recorded at physiological maturity.As indicated previously, in the greenhouse experiment, water restriction was achieved by withholding regular watering. In the field trials, water restriction was performed by reducing the amount of water supplied through irrigation. The phenological stage at which water restriction was imposed was the primary variable considered in both experimental settings.Statistical analysisData was statistically analyzed as indicated in each figure legend using Prism 9 (GraphPad Software Inc., California, USA). Outlier values were detected by the ROUT method and excluded from the analyses. When required, data sets were transformed to meet the statistical test requirements.ResultsIn the greenhouse experiment using the commercial maize hybrid DOW 510 PW, water deficit imposed at V2 negatively affected height and shoot fresh weight in uninoculated plants, with reductions ranging from 20% to 67% compared to full-irrigated plants (<a class="elsevierStyleCrossRef" href="#fig0005">Figs. 1a–c</a>). Notably, inoculation with strain Az19 prevented the adverse effects of water deficit on these growth parameters, whereas Az39 did not (<a class="elsevierStyleCrossRef" href="#fig0005">Figs. 1a–c</a>). Water deficit also decreased the RWC of the last expanded leaf by 34% in the uninoculated control, but inoculated plants maintained the same RWC as irrigated plants, irrespective of the A. argentinense strain supplied (<a class="elsevierStyleCrossRef" href="#fig0005">Fig. 1</a>d).<elsevierMultimedia ident="fig0005"></elsevierMultimedia>With regard to the reproductive performance of the maize plants, none of the uninoculated plants produced ears under water restrictions (0% of fertile plants) (<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>). Inoculation mitigated this effect irrespective of the A. argentinense strain supplied. Compared to uninoculated plants not subjected to water deficits, the grain weight per plant decreased significantly due to water restriction in Az39-inoculated plants but not in Az19-inoculated plants (<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>).<elsevierMultimedia ident="tbl0005"></elsevierMultimedia>Despite the fact that inoculation with Az19 had no significant effect on the full-watered treatments, in water-restricted plants, inoculation with this strain mitigated to a greater extent the negative effect of water limitation on yield components, leading to a mean ear weight and a mean grain weight per plant 2-fold and 5-fold increased, respectively, compared to those recorded for plants inoculated with strain Az39 (<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>).In the first field assay with a commercial hybrid, water restriction imposed at flowering significantly decreased the yield and increased the empty ear rate compared to full-watered controls, irrespective of the inoculation treatments (<a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>). In the Az19 treatment, water restriction at grain filling resulted in no significant yield reductions whereas in Az39, a significant yield decrease occurred (<a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>).<elsevierMultimedia ident="tbl0010"></elsevierMultimedia>A paired sample t-test was used to analyze the incidence of inoculation treatments on grain yield under water restriction at flowering (<a class="elsevierStyleCrossRef" href="#tbl0015">Table 3</a>). Only the grain yield of Az19-inoculated plants subjected to water restriction did not differ significantly from that of the watered control plants (p≤0.01).<elsevierMultimedia ident="tbl0015"></elsevierMultimedia>In the second field assay where a drought-sensitive (GS) and a drought-tolerant (GT) genotype were used, the 3-way ANOVA indicated that the interaction between the factors (water treatment, inoculation, and genotype) was not significant. Subsequent analyses were conducted by 2-way ANOVA taking water and the inoculation treatment as independent variables.Az19 inoculation positively affected the height of plants grown under optimum irrigation conditions, but not under water deficit (<a class="elsevierStyleCrossRef" href="#fig0010">Fig. 2</a>).<elsevierMultimedia ident="fig0010"></elsevierMultimedia>Our data showed that water deficit at flowering significantly increased the empty ear rate in both non-inoculated and Az19-inoculated groups when compared to their respective irrigated controls (<a class="elsevierStyleCrossRef" href="#fig0010">Fig. 2</a>). However, the significance of this increase was higher for non-inoculated plants (p=0.02 and p=0.08 for non-inoculated and Az19-inoculated plants). The yield was significantly reduced by water deficit at flowering in non-inoculated plants, but not in inoculated plants.DiscussionIn nature, plants thriving in arid and semiarid regions usually have evolved adaptive features that enable them to cope with water limitations, in most cases, to the detriment of seed production<a class="elsevierStyleCrossRef" href="#bib0175">3</a>. However, cereals typically cultivated in humid/subhumid zones, such as maize, sugarcane, or wheat, do not possess such mechanisms<a class="elsevierStyleCrossRef" href="#bib0175">3</a>; thus, rhizospheric microorganisms adapted to adverse conditions may partially compensate for such environmental limitations<a class="elsevierStyleCrossRef" href="#bib0270">22</a>.Under controlled conditions, the collection strain A. argentinense Az19 mitigated the negative effects of water deficit to a greater extent than A. argentinense Az39, the most frequent Azospirillum strain included in commercial inoculants in Argentina. This was evidenced by lower reductions in shoot fresh/dry weight, plant height, percentage of fertile plants, and ear weight<a class="elsevierStyleCrossRef" href="#bib0215">11</a>. In maize and other cereals, the percentage of fertile plants, the ear number per plant, the kernel number per ear, and the 1000-kernel weight are key yield components that determine the final grain yield. Therefore, the maximum yield potential of these crops is expected to be determined at flowering<a class="elsevierStyleCrossRef" href="#bib0160">1</a>.It is well-known that water deficit at reproductive stages impairs grain yield<a class="elsevierStyleCrossRef" href="#bib0305">29</a>, but water restrictions at vegetative stages may also compromise yield due to the impact on photosynthetic biomass production<a class="elsevierStyleCrossRef" href="#bib0265">21</a>. In this research, only repeated water restrictions (at V2 and flowering) had adverse consequences on central yield components such as the percentage of fertile plants, in line with data reviewed by Fahad et al.<a class="elsevierStyleCrossRef" href="#bib0200">8</a>. Since both Azospirillum strains, Az39 and Az19, are good in vitro indole acetic acid producers, this differential behavior regarding plant response to water deficit seems not to be directly related to the ability of these microorganisms to stimulate root system development, and other mechanisms may be involved. In this regard, in a previous greenhouse experiment in which the same maize hybrid (DOW 510 PW) was subjected to drought, plants inoculated with Az19 showed higher RWC in the last expanded leaf and higher proline content in the roots than plants inoculated with Az39<a class="elsevierStyleCrossRef" href="#bib0215">11</a>, indicating that a more efficient osmotic adjustment was probably taking place in plants inoculated with the former strain. Other researchers also observed higher proline content in Azospirillum inoculated-maize plants subjected to drought<a class="elsevierStyleCrossRefs" href="#bib0185">5,14</a>, along with changes in specific water status parameters including increased water potential and apoplastic water fraction, and decreased cell wall modulus of elasticity<a class="elsevierStyleCrossRef" href="#bib0190">6</a>. Interestingly, the high in vitro osmotic tolerance of the strain Az19 was related to high trehalose production in pure cultures subjected to osmotic stress<a class="elsevierStyleCrossRef" href="#bib0215">11</a>; this differential osmoadaptive response of strain Az19 over strain Az39 would contribute to improving plant fitness under water limitations.In our field trial, only water deficit at flowering negatively affected plant yield, and Az19 inoculation mitigated the adverse effects caused by this water restriction. Several authors have indicated that water deficit during flowering impairs the reproductive performance of corn<a class="elsevierStyleCrossRefs" href="#bib0180">4,12,26</a>. Our results agree with those of Casanovas et al.<a class="elsevierStyleCrossRef" href="#bib0185">5</a>, who found that inoculation with A. baldaniorum Sp245 resulted in double benefit in corn plants under water deficit by decreasing the number of aborted grains and maintaining the weight of individual grains. In addition, Naseri et al.<a class="elsevierStyleCrossRef" href="#bib0280">24</a> reported that biofertilization with A. argentinense mitigated water deficit damage in field corn trials.In the field trial using different genotypes, we observed that the inoculation effect was not dependent on the genotype, although several authors have already highlighted the importance of plant genotype in responses of cereals to inoculation with PGPR<a class="elsevierStyleCrossRefs" href="#bib0235">15,23</a>. In this sense, García de Salomone and Döbereiner<a class="elsevierStyleCrossRef" href="#bib0210">10</a> attributed differences in maize response to Azospirillum inoculation in field assays carried out in Argentina to the interactions between genotypes and strains. Likewise, in a field trial that comprised 5 wheat genotypes and two Azospirillum strains, an effect of the interaction between both factors was observed for some parameters<a class="elsevierStyleCrossRef" href="#bib0240">16</a>.ConclusionObtaining new strains with improved plant growth-promoting abilities is a permanent challenge. Our results highlight Az19 as a promising PGPR strain, particularly useful to improve maize yield under water deficit at V2 or flowering.Although Az19 and Az39 strains showed similar positive effects on irrigated plants, strain Az19 was more efficient than Az39 in reducing the adverse effects of water limitation on growth and yield parameters in maize.Our results show that inoculation with selected PGPR may mitigate the adverse impact of water scarcity during critical phenological stages, such as flowering and grain filling, on maize yield. The development of biological products based on this microorganism can be available for farmers faster than resistant or tolerant cultivars obtained through plant breeding, a point that acquires particular significance in light of growing food demands and climate change.Conflict of interestNone declared." "textoCompletoSecciones" => array:1 [ "secciones" => array:14 [ 0 => array:3 [ "identificador" => "xres1976944" "titulo" => "Highlights" "secciones" => array:1 [ 0 => array:1 [ "identificador" => "abst0005" ] ] ] 1 => array:3 [ "identificador" => "xres1976946" "titulo" => "Abstract" "secciones" => array:1 [ 0 => array:1 [ "identificador" => "abst0010" ] ] ] 2 => array:2 [ "identificador" => "xpalclavsec1700518" "titulo" => "Keywords" ] 3 => array:2 [ "identificador" => "xpalclavsec1700517" "titulo" => "Abbreviations" ] 4 => array:3 [ "identificador" => "xres1976945" "titulo" => "Resumen" "secciones" => array:1 [ 0 => array:1 [ "identificador" => "abst0015" ] ] ] 5 => array:2 [ "identificador" => "xpalclavsec1700519" "titulo" => "Palabras clave" ] 6 => array:2 [ "identificador" => "sec0005" "titulo" => "Introduction" ] 7 => array:3 [ "identificador" => "sec0010" "titulo" => "Materials and methods" "secciones" => array:5 [ 0 => array:2 [ "identificador" => "sec0015" "titulo" => "Plant material" ] 1 => array:2 [ "identificador" => "sec0020" "titulo" => "Azospirillum argentinense strains used for maize inoculation" ] 2 => array:2 [ "identificador" => "sec0025" "titulo" => "Greenhouse experiment" ] 3 => array:2 [ "identificador" => "sec0030" "titulo" => "Field assays" ] 4 => array:2 [ "identificador" => "sec0035" "titulo" => "Statistical analysis" ] ] ] 8 => array:2 [ "identificador" => "sec0040" "titulo" => "Results" ] 9 => array:2 [ "identificador" => "sec0045" "titulo" => "Discussion" ] 10 => array:2 [ "identificador" => "sec0050" "titulo" => "Conclusion" ] 11 => array:2 [ "identificador" => "sec0055" "titulo" => "Conflict of interest" ] 12 => array:2 [ "identificador" => "xack692869" "titulo" => "Acknowledgments" ] 13 => array:1 [ "titulo" => "References" ] ] ] "pdfFichero" => "main.pdf" "tienePdf" => true "fechaRecibido" => "2022-07-15" "fechaAceptado" => "2023-01-24" "PalabrasClave" => array:2 [ "en" => array:2 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Keywords" "identificador" => "xpalclavsec1700518" "palabras" => array:4 [ 0 => "Drought" 1 => "Phenological stages" 2 => "PGPR" 3 => "Zea mays" ] ] 1 => array:4 [ "clase" => "abr" "titulo" => "Abbreviations" "identificador" => "xpalclavsec1700517" "palabras" => array:6 [ 0 => "CFU" 1 => "EEA" 2 => "PGPR" 3 => "RWC" 4 => "WR-Fl" 5 => "WR-Gf" ] ] ] "es" => array:1 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Palabras clave" "identificador" => "xpalclavsec1700519" "palabras" => array:4 [ 0 => "Estadios fenológicos" 1 => "PGPR" 2 => "Sequía" 3 => "Zea mays" ] ] ] ] "tieneResumen" => true "highlights" => array:2 [ "titulo" => "Highlights" "resumen" => "<ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005">•A. argentinense Az19 mitigated maize growth reduction due to water deficit (WD) at V2.</li><li class="elsevierStyleListItem" id="lsti0010">•WD at flowering reduced maize yield less in Az19- than in Az39-inoculated plants.</li><li class="elsevierStyleListItem" id="lsti0015">•Az19 is a promising PGPR strain to be used as an inoculant in areas subjected to WD.</li></ul>" ] "resumen" => array:2 [ "en" => array:2 [ "titulo" => "Abstract" "resumen" => "Water deficit constitutes a severe limitation to agricultural productivity. In the context of sustainable crop production, the potential of microbial biotechnology to increase plant drought tolerance and improve crop yields under adverse conditions is gaining relevance. This work aimed to compare the performance of Azospirillumargentinense strain Az19 to that of strain Az39, the most widely used for commercial inoculants, when inoculated in maize plants exposed to water deficit. For this purpose, greenhouse and field assays were conducted. In the greenhouse experiment, strain Az19 prevented the adverse effect of water deficit at V2 stage on maize growth. Moreover, the percentage of fertile plants and the ear weight decreased significantly under water deficits imposed at V2 and flowering in Az39-inoculated plants but not in Az19-inoculated plants. In the first field trial with the commercial maize hybrid DOW DS 515 PW, Az19-inoculated plants were those which better tolerated the water deficit imposed. In the second field trial, two maize genotypes with differential drought sensitivity (LP 29×LP 2542, sensitive; LP 882 (923)×LP 4703, tolerant) were tested. Higher tolerance to water deficit was detected in plants inoculated with A. argentinense Az19, with a noticeable effect on grain yield components in the sensitive genotype. Based on these results, we propose the use of A. argentinense Az19 for the formulation of more targeted Azospirillum-based inoculants, suitable for agroecological areas subjected to seasonal water deficits." ] "es" => array:2 [ "titulo" => "Resumen" "resumen" => "El déficit hídrico constituye una severa limitación a la productividad agrícola. En el marco de la producción sostenible de cultivos, la biotecnología microbiana está cobrando relevancia para aumentar la tolerancia a la sequía y mejorar el rendimiento de los cultivos en condiciones adversas. El propósito de este trabajo fue comparar la acción de la cepa de Azospirillum argentinense Az19, con tolerancia in vitro a estresores abióticos, con la cepa Az39, utilizada ampliamente para la formulación de inoculantes comerciales, al inocularlas en plantas sometidas a déficit hídrico. Se realizaron ensayos de invernadero y de campo. En invernadero, la cepa Az19 evitó el impacto adverso del déficit hídrico en el estadio V2 sobre el crecimiento del maíz. Además, el porcentaje de plantas con espigas y el peso de la espiga disminuyó significativamente con la restricción hídrica aplicada en V2 y en floración en plantas inoculadas con la cepa Az39, pero no en las inoculadas con Az19. En el primer ensayo de campo con el maíz híbrido comercial DOW DS 515 PW las plantas inoculadas con Az19 fueron las que mejor toleraron la deficiencia hídrica. En el segundo ensayo de campo se utilizaron dos genotipos de maíz con sensibilidad diferencial a la sequía. La inoculación con Az19 condujo a una mayor tolerancia al déficit hídrico, con un efecto detectable en algunos componentes del rendimiento en el genotipo sensible. Sobre la base de estos resultados, proponemos el empleo de A. argentinense Az19 para la formulación de inoculantes basados en Azospirillum especialmente indicados para áreas agroecológicas que experimenten períodos de déficit hídrico." ] ] "multimedia" => array:6 [ 0 => array:7 [ "identificador" => "fig0005" "etiqueta" => "Figure 1" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr1.jpeg" "Alto" => 1867 "Ancho" => 3175 "Tamanyo" => 205129 ] ] "descripcion" => array:1 [ "en" => "Effect of inoculation and water deficit at V2 on maize growth and physiological parameters (greenhouse experiment). Mean values and standard deviation determined at V3 are shown (n=12). (a) Plant height; (b) shoot fresh weight; (c) shoot dry weight; (d) relative water content of the last expanded leaf. Control: no water deficit; WR: water restriction; NI: non-inoculated. Data was square root-transformed and analyzed by two-way ANOVA plus the Tukey's test. Asterisks indicate significant differences between control and WR groups within each inoculation treatment (p˂0.1)." ] ] 1 => array:7 [ "identificador" => "fig0010" "etiqueta" => "Figure 2" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr2.jpeg" "Alto" => 959 "Ancho" => 3175 "Tamanyo" => 268312 ] ] "descripcion" => array:1 [ "en" => "Changes in maize height (A), percentage of empty ears (B), and yield (C) according to irrigation and inoculation treatments (field assay 2). Mean values and standard deviation determined at physiological maturity are shown. Sampling was performed as described in “Materials and methods” section. C: full-irrigated control; WR-Fl: water restriction at flowering; WR-Gf: water restriction at grain filling; GS: drought-sensitive genotype; GT: drought-tolerant genotype. Data was square root-transformed (B) or not transformed (A and C) and analyzed by two-way ANOVA plus the Tukey's test. Different letters indicate significant differences between treatments irrespective of the plant genotype (p˂0.1)." ] ] 2 => array:8 [ "identificador" => "tbl0005" "etiqueta" => "Table 1" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at1" "detalle" => "Table " "rol" => "short" ] ] "tabla" => array:2 [ "leyenda" => "Mean values obtained at physiological maturity are shown. C: irrigated on demand; WR: water restriction; Ni: not inoculated. Asterisks indicate significant differences between C and WR groups within each inoculation treatment, according to two-way ANOVA followed by Šídák's multiple range test (p≤0.1)." "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Treatment \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Fertile plants (%) \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Ear weight (g) \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Grain weight per plant (g) \t\t\t\t\t\t\n \t\t\t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">C:Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">39.3±12.34 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">10.65±5.07 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5.2±2.12 \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">C:Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">33.0±14.75 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">8.06±4.28 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">3.66±2.26 \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">C:Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">39.6±6.6 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">12.73±3.69 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5.69±2.11 \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR:Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0* \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">– \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">– \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR:Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">33.2±18.75 ns \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5.62±1.66 ns \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">2.35±0.02 ns \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR:Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">13.2±8.08 ns \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">2.94±1.86* \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0.44±0.43* \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab3290888.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Effect of inoculation and water deficit imposed at V2 and flowering on maize productivity parameters (greenhouse experiment)." ] ] 3 => array:8 [ "identificador" => "tbl0010" "etiqueta" => "Table 2" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at2" "detalle" => "Table " "rol" => "short" ] ] "tabla" => array:2 [ "leyenda" => "Mean values obtained at physiological maturity are shown. Control: full-irrigated; WR-Fl: water restriction at flowering; WR-Gf: water restriction at grain filling; NI: non-inoculated. Different letters indicate significant differences between different water treatments within each inoculation group according to two-way ANOVA followed by Tukey's multiple range test (p˂0.1)." "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Water treatment \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Inoculation treatment \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " colspan="2" align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Yield (kg/ha)</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " colspan="2" align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Empty ear rate (%)</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Control \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">9338±634 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">4.3±1.52 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">4564±1007 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">29.3±13.35 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Gf \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">7465±942 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">1.6±1.18 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Control \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">10187±697 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">2.3±1.41 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5782±1009 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">14.9±7.09 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Gf \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">8122±682 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">AB \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Control \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">10060±991 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">1.7±1.25 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5095±770 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">20.4±8.45 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Gf \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">8028±695 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">B \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">2.6±1.63 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">A \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab3290886.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Effect of inoculation and water restriction on maize productivity parameters (field assay 1)." ] ] 4 => array:8 [ "identificador" => "tbl0015" "etiqueta" => "Table 3" "tipo" => "MULTIMEDIATABLA" "mostrarFloat" => true "mostrarDisplay" => false "detalles" => array:1 [ 0 => array:3 [ "identificador" => "at3" "detalle" => "Table " "rol" => "short" ] ] "tabla" => array:2 [ "leyenda" => "Ni: non-inoculated; WR-Fl: plants subjected to water restriction at flowering. Asterisks indicate significant differences at p˂0.01." "tablatextoimagen" => array:1 [ 0 => array:2 [ "tabla" => array:1 [ 0 => """ <table border="0" frame="\n \t\t\t\t\tvoid\n \t\t\t\t" class=""><thead title="thead"><tr title="table-row"><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Group 1 \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Group 2 \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Mean(1) \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Mean(2) \t\t\t\t\t\t\n \t\t\t\t\t\t</th><th class="td" title="\n \t\t\t\t\ttable-head\n \t\t\t\t " align="center" valign="\n \t\t\t\t\ttop\n \t\t\t\t" scope="col" style="border-bottom: 2px solid black">p-Value \t\t\t\t\t\t\n \t\t\t\t\t\t</th></tr></thead><tbody title="tbody"><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl: Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Watered Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">4564 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">9338 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0.004** \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl: Az19 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Watered Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5782 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">9338 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0.018 \t\t\t\t\t\t\n \t\t\t\t</td></tr><tr title="table-row"><td class="td-with-role" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">WR-Fl: Az39 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="left" valign="\n \t\t\t\t\ttop\n \t\t\t\t">Watered Ni \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">5095 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">9338 \t\t\t\t\t\t\n \t\t\t\t</td><td class="td" title="\n \t\t\t\t\ttable-entry\n \t\t\t\t " align="char" valign="\n \t\t\t\t\ttop\n \t\t\t\t">0.003** \t\t\t\t\t\t\n \t\t\t\t</td></tr></tbody></table> """ ] "imagenFichero" => array:1 [ 0 => "xTab3290887.png" ] ] ] ] "descripcion" => array:1 [ "en" => "Paired sample t-test for grain yield (field assay 1)." ] ] 5 => array:5 [ "identificador" => "eq0005" "tipo" => "MULTIMEDIAFORMULA" "mostrarFloat" => false "mostrarDisplay" => true "Formula" => array:5 [ "Matematica" => "RWC⁡ (%)=(fresh weight−dry weight)×100turgor weight−dry weight." 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Original article

La inoculación con Azospirillum argentinense Az19 mejora el rendimiento del maíz expuesto a déficit hídrico durante estadios clave del desarrollo vegetal

Julia Elena Garcíaa, Mónica Ruizb, Guillermo Andrés Maronichec,d, Cecilia Creusc, Mariana Puentea, Myriam Sara Zawoznike,

Corresponding author

myriamz@ffyb.uba.ar

Corresponding author.

, María Daniela Groppaf

a Instituto Nacional de Tecnología Agropecuaria (INTA), Instituto de Microbiología y Zoología Agrícola, Argentina

b Instituto Nacional de Tecnología Agropecuaria (INTA), Estación Experimental San Juan, Argentina

c Facultad de Ciencias Agrarias, Universidad Nacional de Mar del Plata, Argentina

d Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina

e Universidad de Buenos Aires, Facultad de Farmacia y Bioquímica, Buenos Aires, Argentina

f Universidad de Buenos Aires, Instituto de Química y Fisicoquímica Biológicas (IQUIFIB), Facultad de Farmacia y Bioquímica, Buenos Aires, Argentina

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        "titulo" => "La inoculaci&#243;n con <span class="elsevierStyleItalic">Azospirillum argentinense</span> Az19 mejora el rendimiento del ma&#237;z expuesto a d&#233;ficit h&#237;drico durante estadios clave del desarrollo vegetal"
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          "en" => "<p id="spar0025" class="elsevierStyleSimplePara elsevierViewall">Changes in maize height &#40;A&#41;&#44; percentage of empty ears &#40;B&#41;&#44; and yield &#40;C&#41; according to irrigation and inoculation treatments &#40;field assay 2&#41;&#46; Mean values and standard deviation determined at physiological maturity are shown&#46; Sampling was performed as described in &#8220;Materials and methods&#8221; section&#46; C&#58; full-irrigated control&#59; WR-Fl&#58; water restriction at flowering&#59; WR-Gf&#58; water restriction at grain filling&#59; GS&#58; drought-sensitive genotype&#59; GT&#58; drought-tolerant genotype&#46; Data was square root-transformed &#40;B&#41; or not transformed &#40;A and C&#41; and analyzed by two-way ANOVA plus the Tukey&#39;s test&#46; Different letters indicate significant differences between treatments irrespective of the plant genotype &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#706;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p>"
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    "textoCompleto" => "<span class="elsevierStyleSections"><span id="sec0005" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0035">Introduction</span><p id="par0020" class="elsevierStylePara elsevierViewall">In Argentina&#44; maize &#40;<span class="elsevierStyleItalic">Zea mays</span> L&#46;&#41; was traditionally cultivated in the Undulating Pampa region &#40;Pampa Ondulada&#41; &#40;32&#176; to 35&#46;8&#176; S and 58&#176; to 62&#176; W&#41;&#44; a temperate-humid region without climatic constraints and adequate for extensive agriculture&#46; However&#44; due to the massive cultivation of soybean in the last decades&#44; maize production has been expanded outside that optimal zone to semiarid environments &#40;&#60;700<span class="elsevierStyleHsp" style=""></span>mm precipitation per year&#41;&#44; where high temperatures and water restrictions are relatively frequent events<a class="elsevierStyleCrossRef" href="#bib0260"><span class="elsevierStyleSup">20</span></a>&#46;</p><p id="par0025" class="elsevierStylePara elsevierViewall">Flowering is a key phenological stage for cereal production&#44; and water deficit occurring during that stage adversely impacts yield due to its negative effect on grain development&#46; Water limitation at flowering leads to impaired reproductive performance&#44; resulting in a decreased number of grains per hectare and decreased final grain weight<a class="elsevierStyleCrossRef" href="#bib0255"><span class="elsevierStyleSup">19</span></a>&#46;</p><p id="par0030" class="elsevierStylePara elsevierViewall">In addition to plant breeding techniques&#44; microbial biotechnology is increasingly positioned as a valuable strategy to improve crop performance under adverse conditions&#46; In this context&#44; the introduction of plant growth-promoting rhizobacteria &#40;PGPR&#41; is an environmental-friendly alternative to chemical treatments for sustainable production systems&#46; PGPR comprise those microorganisms that colonize the rhizosphere of plants and promote plant growth&#46; Within this group&#44; the genus <span class="elsevierStyleItalic">Azospirillum</span> has been one of the most studied and is usually regarded as a model bacterium to study plant-bacteria interactions<a class="elsevierStyleCrossRef" href="#bib0295"><span class="elsevierStyleSup">27</span></a>&#46; <span class="elsevierStyleItalic">Azospirillum</span> can interact with a wide range of plant species due to its recognized metabolic versatility<a class="elsevierStyleCrossRef" href="#bib0295"><span class="elsevierStyleSup">27</span></a>&#46; For example&#44; the strain Az39 of <span class="elsevierStyleItalic">Azospirillum argentinense</span>&#44; used in most commercial formulations in Argentina&#44; was tested in various cereals and was found to increase yield significantly<a class="elsevierStyleCrossRefs" href="#bib0195"><span class="elsevierStyleSup">7&#44;9&#44;30</span></a>&#46;</p><p id="par0035" class="elsevierStylePara elsevierViewall">It has also been documented that plants inoculated with PGPR exhibit greater tolerance to abiotic stresses such as hydric and osmotic stresses<a class="elsevierStyleCrossRef" href="#bib0250"><span class="elsevierStyleSup">18</span></a>&#46; The beneficial effects of PGPR inoculation in plants subjected to osmotic stress include biomass increase and water status improvement<a class="elsevierStyleCrossRef" href="#bib0245"><span class="elsevierStyleSup">17</span></a>&#46; Under field conditions&#44; wheat inoculation with <span class="elsevierStyleItalic">A&#46; baldaniorum</span> Sp245 increased cell wall elasticity in the flag leaf and reduced yield losses due to moderate drought at the anthesis stage&#46; In addition&#44; this strain mitigated the negative effect of drought and led to increased grain yield and mineral content &#40;Mg&#44; K&#44; and Ca&#41; compared to non-inoculated plants<a class="elsevierStyleCrossRef" href="#bib0190"><span class="elsevierStyleSup">6</span></a>&#46;</p><p id="par0040" class="elsevierStylePara elsevierViewall">Previously&#44; we characterized the strain Az19 of <span class="elsevierStyleItalic">A&#46; argentinense</span> from the PGPR collection of IMyZA INTA and observed that this strain had similar PGPR features but better attributes to cope with osmotic stress than <span class="elsevierStyleItalic">A&#46; argentinense</span> Az39&#44; the strain typically included in commercial inoculants<a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">11</span></a>&#46; Az19 is a collection strain isolated in 1982 from wheat field fallow&#46; Since this strain has already demonstrated higher <span class="elsevierStyleItalic">in vitro</span> survival under saline and osmotic stress than Az39 and higher trehalose production<a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">11</span></a>&#44; we hypothesized that this bacterium could also display higher survival in the soil&#47;root interface and thus contribute to sustain plant growth and yield under water deficit&#44; which usually entails osmotic imbalances&#46; The purpose of this study was to assess if seed inoculation with the strain Az19 ameliorates the negative effects of water restriction at specific phenological stages on the yield of a commercial maize hybrid to a greater extent than the Az39 strain &#40;greenhouse experiment and field assay 1&#41; and on different contrasting maize genotypes for their sensitivity to drought &#40;field assay 2&#41;&#46;</p></span><span id="sec0010" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0040">Materials and methods</span><span id="sec0015" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0045">Plant material</span><p id="par0045" class="elsevierStylePara elsevierViewall">Two commercial maize hybrids&#44; DOW 510 PW &#40;greenhouse experiment&#41; and DOW DS 515 PW &#40;field experiment 1&#41;&#44; and two maize genotypes obtained at INTA Pergamino&#44; Argentina&#44; LP 29<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 2542 &#40;drought-sensitive&#41; and LP 882 &#40;923&#41;<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 4703 &#40;drought-tolerant&#41; &#40;field experiment 2&#41;&#44; were used&#46;</p></span><span id="sec0020" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0050"><span class="elsevierStyleItalic">Azospirillum argentinense</span> strains used for maize inoculation</span><p id="par0050" class="elsevierStylePara elsevierViewall">Az39 and Az19 were obtained from the PGPR collection of IMyZA INTA Castelar&#46; Az39 was used as the reference strain&#46;</p><p id="par0055" class="elsevierStylePara elsevierViewall">Both <span class="elsevierStyleItalic">A&#46; brasilense</span> strains were cultured in OAB liquid medium<a class="elsevierStyleCrossRef" href="#bib0285"><span class="elsevierStyleSup">25</span></a> supplemented with 1<span class="elsevierStyleHsp" style=""></span>g&#47;l NH<span class="elsevierStyleInf">4</span>Cl &#40;for an optimal bacterial growth&#41; at 30<span class="elsevierStyleHsp" style=""></span>&#176;C in a rotary shaker at 180<span class="elsevierStyleHsp" style=""></span>rpm for 48<span class="elsevierStyleHsp" style=""></span>h&#46; Viable cell number &#40;CFU&#47;ml&#41; was quantified using the drop plate method<a class="elsevierStyleCrossRef" href="#bib0225"><span class="elsevierStyleSup">13</span></a> on plates containing Congo Red medium<a class="elsevierStyleCrossRef" href="#bib0300"><span class="elsevierStyleSup">28</span></a>&#46;</p></span><span id="sec0025" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0055">Greenhouse experiment</span><p id="par0060" class="elsevierStylePara elsevierViewall">Surface disinfected &#40;ethanol 80&#37; and sodium hypochlorite 4&#37;&#44; v&#47;v&#41; seeds of the maize hybrid DOW 510 PW were pregerminated in a wet chamber at 25<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; and uniform-developed seedlings were inoculated 24<span class="elsevierStyleHsp" style=""></span>h later by immersion for 2<span class="elsevierStyleHsp" style=""></span>h in an inoculum containing 5<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>10<span class="elsevierStyleSup">7</span> colony-forming units &#40;CFU&#41; of the corresponding <span class="elsevierStyleItalic">A&#46; argentinense</span> strain per seedling&#46; A set of non-inoculated seedlings was included&#46; Emerging seedlings were subsequently transferred to 10<span class="elsevierStyleHsp" style=""></span>l plastic pots containing a mixture of sterile sand&#44; soil&#44; vermiculite&#44; and perlite &#40;3&#58;3&#58;3&#58;1&#41; as substrate&#46;</p><p id="par0065" class="elsevierStylePara elsevierViewall">All treatments &#40;including controls&#41; were irrigated with Hoagland nutrient solution&#46; Water restriction was applied once at V2 or twice at V2 and flowering&#44; and plants were harvested at V3 or physiological maturity&#44; respectively&#46; Water deficit at V2 was achieved by withholding the irrigation for 10 days while water deficit at flowering was achieved by withholding irrigation for 20 days after flowering&#46; Male and female flowering dates were recorded when 50&#37; of the plants were shedding pollen or showed 1-cm long stigmas&#44; respectively&#46;</p><p id="par0070" class="elsevierStylePara elsevierViewall">A randomized complete block design consisting of 3 blocks with 12 plants per treatment was used&#46; Length&#44; fresh weight&#44; and dry weight of the aerial part and the relative water content &#40;RWC&#41; of the last expanded leaf were measured at V3 &#40;plants exposed to water deficit at V2&#41;&#46; RWC was determined according to the method described by Barrs and Weatherley<a class="elsevierStyleCrossRef" href="#bib0170"><span class="elsevierStyleSup">2</span></a> based on the following formula&#58;<elsevierMultimedia ident="eq0005"></elsevierMultimedia></p><p id="par0075" class="elsevierStylePara elsevierViewall">At physiological maturity&#44; the percentage of fertile plants &#40;grain-producing plants&#41;&#44; the ear weight&#44; and the grain weight per plant were determined &#40;plants exposed to water deficit at V2 and flowering&#41;&#46;</p></span><span id="sec0030" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0060">Field assays</span><p id="par0080" class="elsevierStylePara elsevierViewall">Both field assays were conducted in plots belonging to the agricultural experimental station of the Instituto Nacional de Tecnolog&#237;a Agropecuaria in San Juan province &#40;INTA EEA San Juan&#41;&#44; located near Pocito city &#40;31&#176;39&#8242;0&#8243;S&#44; 68&#176;33&#8242;0&#8243;W&#41;&#44; during the maize campaign 2015&#47;16&#46; We selected this location because of the recognized scarcity of rainfall&#44; which is appropriate to mimic a water-deficient condition requiring controlled irrigation&#46;</p><p id="par0085" class="elsevierStylePara elsevierViewall">The soil of this area is classified as Entisol&#44; and since 2009&#44; these plots have been under maize production&#44; with fallow between cycles&#46; Plots received 200<span class="elsevierStyleHsp" style=""></span>kg N&#47;ha &#40;100<span class="elsevierStyleHsp" style=""></span>kg at sowing and 100<span class="elsevierStyleHsp" style=""></span>kg at V7&#41;&#46; The mean temperature at silking &#40;&#177;15 days&#41; was 30&#46;2<span class="elsevierStyleHsp" style=""></span>&#176;C&#44; and from that stage to grain filling was 24&#46;1<span class="elsevierStyleHsp" style=""></span>&#176;C&#46; Accumulated rainfall in the same periods was 13&#46;5 and 7&#46;2<span class="elsevierStyleHsp" style=""></span>mm&#44; respectively&#46;</p><p id="par0090" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">Field assay 1</span>&#46; The commercial maize hybrid DOW DS 515 PW was used&#46; Before planting&#44; seeds were inoculated with the corresponding <span class="elsevierStyleItalic">A&#46; argentinense</span> strain &#40;12<span class="elsevierStyleHsp" style=""></span>ml&#47;kg seeds&#59; 1<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>10<span class="elsevierStyleSup">9</span><span class="elsevierStyleHsp" style=""></span>CFU&#47;ml of inoculant&#41; and then sown at a plant density of 70<span class="elsevierStyleHsp" style=""></span>000<span class="elsevierStyleHsp" style=""></span>plants&#47;ha&#44; with a distance of 0&#46;63<span class="elsevierStyleHsp" style=""></span>m between rows and 0&#46;19<span class="elsevierStyleHsp" style=""></span>m between plants&#46; A split-plot experimental design was used&#44; the main plots corresponding to factor A&#44; water restriction&#44; and the secondary plots to factor B&#44; inoculation&#44; with 5 repetitions &#40;plots&#41; per treatment&#46; Each plot included eight 5-m long rows&#46;</p><p id="par0095" class="elsevierStylePara elsevierViewall">Water restriction &#40;factor A&#41; included a full-irrigated control&#44; irrigated at 100&#37; of the calculated evapotranspiration &#40;ETc&#41; throughout the entire crop cycle&#44; and two alternative water deficit treatments&#58; one imposed at flowering&#44; with the irrigation reduced to 25&#37; of the ETc between &#8722;15 to &#43;15 days respect to the flowering date &#40;WR-Fl&#41;&#44; and the other one imposed at grain filling&#44; with the irrigation reduced to 25&#37; of the ETc from grain set up to physiological maturity &#40;WR-Gf&#41;&#46; Inoculation treatments &#40;factor B&#41; were the same as in the greenhouse experiment&#58; Az39&#44; Az19&#44; none &#40;control&#41;&#46;</p><p id="par0100" class="elsevierStylePara elsevierViewall">Male and female flowering dates were recorded when more than 15 plants per row were shedding pollen and had 1-cm long stigmas&#44; respectively&#46; Irrigation was performed <span class="elsevierStyleItalic">via</span> a surface drip irrigation system with hoses of 1&#46;27<span class="elsevierStyleHsp" style=""></span>cm in diameter and emitters every 25<span class="elsevierStyleHsp" style=""></span>cm&#44; with a flow of 2&#46;5<span class="elsevierStyleHsp" style=""></span>l&#47;h&#46; The irrigation rate was established based on the calculated crop evapotranspiration&#46; Yield &#40;kg&#47;ha&#41; and empty ear rates &#40;&#37;&#41; were evaluated at physiological maturity by sampling two random rows&#46; The plant stand was uniform across treatments&#46;</p><p id="par0105" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">Field assay 2&#46;</span> Only strain Az19 was used as inoculation treatment&#44; and two non-commercial hybrids provided by the Breeding Program of INTA EEA Pergamino&#44; Argentina&#44; were tested in this trial&#58; the drought-sensitive genotype LP 29<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 2542 and the drought-tolerant genotype LP 882 &#40;923&#41;<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 4703&#46; Water restriction was imposed as in field assay 1&#44; and yield and empty ear rate were evaluated as described&#46; Plant height was an additional parameter recorded at physiological maturity&#46;</p><p id="par0110" class="elsevierStylePara elsevierViewall">As indicated previously&#44; in the greenhouse experiment&#44; water restriction was achieved by withholding regular watering&#46; In the field trials&#44; water restriction was performed by reducing the amount of water supplied through irrigation&#46; The phenological stage at which water restriction was imposed was the primary variable considered in both experimental settings&#46;</p></span><span id="sec0035" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0065">Statistical analysis</span><p id="par0115" class="elsevierStylePara elsevierViewall">Data was statistically analyzed as indicated in each figure legend using Prism 9 &#40;GraphPad Software Inc&#46;&#44; California&#44; USA&#41;&#46; Outlier values were detected by the ROUT method and excluded from the analyses&#46; When required&#44; data sets were transformed to meet the statistical test requirements&#46;</p></span></span><span id="sec0040" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0070">Results</span><p id="par0120" class="elsevierStylePara elsevierViewall">In the greenhouse experiment using the commercial maize hybrid DOW 510 PW&#44; water deficit imposed at V2 negatively affected height and shoot fresh weight in uninoculated plants&#44; with reductions ranging from 20&#37; to 67&#37; compared to full-irrigated plants &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Figs&#46; 1a&#8211;c</a>&#41;&#46; Notably&#44; inoculation with strain Az19 prevented the adverse effects of water deficit on these growth parameters&#44; whereas Az39 did not &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Figs&#46; 1a&#8211;c</a>&#41;&#46; Water deficit also decreased the RWC of the last expanded leaf by 34&#37; in the uninoculated control&#44; but inoculated plants maintained the same RWC as irrigated plants&#44; irrespective of the <span class="elsevierStyleItalic">A&#46; argentinense</span> strain supplied &#40;<a class="elsevierStyleCrossRef" href="#fig0005">Fig&#46; 1</a>d&#41;&#46;</p><elsevierMultimedia ident="fig0005"></elsevierMultimedia><p id="par0125" class="elsevierStylePara elsevierViewall">With regard to the reproductive performance of the maize plants&#44; none of the uninoculated plants produced ears under water restrictions &#40;0&#37; of fertile plants&#41; &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#41;&#46; Inoculation mitigated this effect irrespective of the <span class="elsevierStyleItalic">A&#46; argentinense</span> strain supplied&#46; Compared to uninoculated plants not subjected to water deficits&#44; the grain weight per plant decreased significantly due to water restriction in Az39-inoculated plants but not in Az19-inoculated plants &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#41;&#46;</p><elsevierMultimedia ident="tbl0005"></elsevierMultimedia><p id="par0130" class="elsevierStylePara elsevierViewall">Despite the fact that inoculation with Az19 had no significant effect on the full-watered treatments&#44; in water-restricted plants&#44; inoculation with this strain mitigated to a greater extent the negative effect of water limitation on yield components&#44; leading to a mean ear weight and a mean grain weight per plant 2-fold and 5-fold increased&#44; respectively&#44; compared to those recorded for plants inoculated with strain Az39 &#40;<a class="elsevierStyleCrossRef" href="#tbl0005">Table 1</a>&#41;&#46;</p><p id="par0135" class="elsevierStylePara elsevierViewall">In the first field assay with a commercial hybrid&#44; water restriction imposed at flowering significantly decreased the yield and increased the empty ear rate compared to full-watered controls&#44; irrespective of the inoculation treatments &#40;<a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>&#41;&#46; In the Az19 treatment&#44; water restriction at grain filling resulted in no significant yield reductions whereas in Az39&#44; a significant yield decrease occurred &#40;<a class="elsevierStyleCrossRef" href="#tbl0010">Table 2</a>&#41;&#46;</p><elsevierMultimedia ident="tbl0010"></elsevierMultimedia><p id="par0140" class="elsevierStylePara elsevierViewall">A paired sample <span class="elsevierStyleItalic">t</span>-test was used to analyze the incidence of inoculation treatments on grain yield under water restriction at flowering &#40;<a class="elsevierStyleCrossRef" href="#tbl0015">Table 3</a>&#41;&#46; Only the grain yield of Az19-inoculated plants subjected to water restriction did not differ significantly from that of the watered control plants &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#8804;<span class="elsevierStyleHsp" style=""></span>0&#46;01&#41;&#46;</p><elsevierMultimedia ident="tbl0015"></elsevierMultimedia><p id="par0145" class="elsevierStylePara elsevierViewall">In the second field assay where a drought-sensitive &#40;GS&#41; and a drought-tolerant &#40;GT&#41; genotype were used&#44; the 3-way ANOVA indicated that the interaction between the factors &#40;water treatment&#44; inoculation&#44; and genotype&#41; was not significant&#46; Subsequent analyses were conducted by 2-way ANOVA taking water and the inoculation treatment as independent variables&#46;</p><p id="par0150" class="elsevierStylePara elsevierViewall">Az19 inoculation positively affected the height of plants grown under optimum irrigation conditions&#44; but not under water deficit &#40;<a class="elsevierStyleCrossRef" href="#fig0010">Fig&#46; 2</a>&#41;&#46;</p><elsevierMultimedia ident="fig0010"></elsevierMultimedia><p id="par0155" class="elsevierStylePara elsevierViewall">Our data showed that water deficit at flowering significantly increased the empty ear rate in both non-inoculated and Az19-inoculated groups when compared to their respective irrigated controls &#40;<a class="elsevierStyleCrossRef" href="#fig0010">Fig&#46; 2</a>&#41;&#46; However&#44; the significance of this increase was higher for non-inoculated plants &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;02 and <span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>0&#46;08 for non-inoculated and Az19-inoculated plants&#41;&#46; The yield was significantly reduced by water deficit at flowering in non-inoculated plants&#44; but not in inoculated plants&#46;</p></span><span id="sec0045" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0075">Discussion</span><p id="par0160" class="elsevierStylePara elsevierViewall">In nature&#44; plants thriving in arid and semiarid regions usually have evolved adaptive features that enable them to cope with water limitations&#44; in most cases&#44; to the detriment of seed production<a class="elsevierStyleCrossRef" href="#bib0175"><span class="elsevierStyleSup">3</span></a>&#46; However&#44; cereals typically cultivated in humid&#47;subhumid zones&#44; such as maize&#44; sugarcane&#44; or wheat&#44; do not possess such mechanisms<a class="elsevierStyleCrossRef" href="#bib0175"><span class="elsevierStyleSup">3</span></a>&#59; thus&#44; rhizospheric microorganisms adapted to adverse conditions may partially compensate for such environmental limitations<a class="elsevierStyleCrossRef" href="#bib0270"><span class="elsevierStyleSup">22</span></a>&#46;</p><p id="par0165" class="elsevierStylePara elsevierViewall">Under controlled conditions&#44; the collection strain <span class="elsevierStyleItalic">A&#46; argentinense</span> Az19 mitigated the negative effects of water deficit to a greater extent than <span class="elsevierStyleItalic">A&#46; argentinense</span> Az39&#44; the most frequent <span class="elsevierStyleItalic">Azospirillum</span> strain included in commercial inoculants in Argentina&#46; This was evidenced by lower reductions in shoot fresh&#47;dry weight&#44; plant height&#44; percentage of fertile plants&#44; and ear weight<a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">11</span></a>&#46; In maize and other cereals&#44; the percentage of fertile plants&#44; the ear number per plant&#44; the kernel number per ear&#44; and the 1000-kernel weight are key yield components that determine the final grain yield&#46; Therefore&#44; the maximum yield potential of these crops is expected to be determined at flowering<a class="elsevierStyleCrossRef" href="#bib0160"><span class="elsevierStyleSup">1</span></a>&#46;</p><p id="par0170" class="elsevierStylePara elsevierViewall">It is well-known that water deficit at reproductive stages impairs grain yield<a class="elsevierStyleCrossRef" href="#bib0305"><span class="elsevierStyleSup">29</span></a>&#44; but water restrictions at vegetative stages may also compromise yield due to the impact on photosynthetic biomass production<a class="elsevierStyleCrossRef" href="#bib0265"><span class="elsevierStyleSup">21</span></a>&#46; In this research&#44; only repeated water restrictions &#40;at V2 and flowering&#41; had adverse consequences on central yield components such as the percentage of fertile plants&#44; in line with data reviewed by Fahad et al&#46;<a class="elsevierStyleCrossRef" href="#bib0200"><span class="elsevierStyleSup">8</span></a>&#46; Since both <span class="elsevierStyleItalic">Azospirillum</span> strains&#44; Az39 and Az19&#44; are good <span class="elsevierStyleItalic">in vitro</span> indole acetic acid producers&#44; this differential behavior regarding plant response to water deficit seems not to be directly related to the ability of these microorganisms to stimulate root system development&#44; and other mechanisms may be involved&#46; In this regard&#44; in a previous greenhouse experiment in which the same maize hybrid &#40;DOW 510 PW&#41; was subjected to drought&#44; plants inoculated with Az19 showed higher RWC in the last expanded leaf and higher proline content in the roots than plants inoculated with Az39<a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">11</span></a>&#44; indicating that a more efficient osmotic adjustment was probably taking place in plants inoculated with the former strain&#46; Other researchers also observed higher proline content in <span class="elsevierStyleItalic">Azospirillum</span> inoculated-maize plants subjected to drought<a class="elsevierStyleCrossRefs" href="#bib0185"><span class="elsevierStyleSup">5&#44;14</span></a>&#44; along with changes in specific water status parameters including increased water potential and apoplastic water fraction&#44; and decreased cell wall modulus of elasticity<a class="elsevierStyleCrossRef" href="#bib0190"><span class="elsevierStyleSup">6</span></a>&#46; Interestingly&#44; the high <span class="elsevierStyleItalic">in vitro</span> osmotic tolerance of the strain Az19 was related to high trehalose production in pure cultures subjected to osmotic stress<a class="elsevierStyleCrossRef" href="#bib0215"><span class="elsevierStyleSup">11</span></a>&#59; this differential osmoadaptive response of strain Az19 over strain Az39 would contribute to improving plant fitness under water limitations&#46;</p><p id="par0175" class="elsevierStylePara elsevierViewall">In our field trial&#44; only water deficit at flowering negatively affected plant yield&#44; and Az19 inoculation mitigated the adverse effects caused by this water restriction&#46; Several authors have indicated that water deficit during flowering impairs the reproductive performance of corn<a class="elsevierStyleCrossRefs" href="#bib0180"><span class="elsevierStyleSup">4&#44;12&#44;26</span></a>&#46; Our results agree with those of Casanovas et al&#46;<a class="elsevierStyleCrossRef" href="#bib0185"><span class="elsevierStyleSup">5</span></a>&#44; who found that inoculation with <span class="elsevierStyleItalic">A&#46; baldaniorum</span> Sp245 resulted in double benefit in corn plants under water deficit by decreasing the number of aborted grains and maintaining the weight of individual grains&#46; In addition&#44; Naseri et al&#46;<a class="elsevierStyleCrossRef" href="#bib0280"><span class="elsevierStyleSup">24</span></a> reported that biofertilization with <span class="elsevierStyleItalic">A&#46; argentinense</span> mitigated water deficit damage in field corn trials&#46;</p><p id="par0180" class="elsevierStylePara elsevierViewall">In the field trial using different genotypes&#44; we observed that the inoculation effect was not dependent on the genotype&#44; although several authors have already highlighted the importance of plant genotype in responses of cereals to inoculation with PGPR<a class="elsevierStyleCrossRefs" href="#bib0235"><span class="elsevierStyleSup">15&#44;23</span></a>&#46; In this sense&#44; Garc&#237;a de Salomone and D&#246;bereiner<a class="elsevierStyleCrossRef" href="#bib0210"><span class="elsevierStyleSup">10</span></a> attributed differences in maize response to <span class="elsevierStyleItalic">Azospirillum</span> inoculation in field assays carried out in Argentina to the interactions between genotypes and strains&#46; Likewise&#44; in a field trial that comprised 5 wheat genotypes and two <span class="elsevierStyleItalic">Azospirillum</span> strains&#44; an effect of the interaction between both factors was observed for some parameters<a class="elsevierStyleCrossRef" href="#bib0240"><span class="elsevierStyleSup">16</span></a>&#46;</p></span><span id="sec0050" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0080">Conclusion</span><p id="par0185" class="elsevierStylePara elsevierViewall">Obtaining new strains with improved plant growth-promoting abilities is a permanent challenge&#46; Our results highlight Az19 as a promising PGPR strain&#44; particularly useful to improve maize yield under water deficit at V2 or flowering&#46;</p><p id="par0190" class="elsevierStylePara elsevierViewall">Although Az19 and Az39 strains showed similar positive effects on irrigated plants&#44; strain Az19 was more efficient than Az39 in reducing the adverse effects of water limitation on growth and yield parameters in maize&#46;</p><p id="par0195" class="elsevierStylePara elsevierViewall">Our results show that inoculation with selected PGPR may mitigate the adverse impact of water scarcity during critical phenological stages&#44; such as flowering and grain filling&#44; on maize yield&#46; The development of biological products based on this microorganism can be available for farmers faster than resistant or tolerant cultivars obtained through plant breeding&#44; a point that acquires particular significance in light of growing food demands and climate change&#46;</p></span><span id="sec0055" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0085">Conflict of interest</span><p id="par0200" class="elsevierStylePara elsevierViewall">None declared&#46;</p></span></span>"
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              "titulo" => "Azospirillum argentinense strains used for maize inoculation"
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              "titulo" => "Greenhouse experiment"
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            3 => "<span class="elsevierStyleItalic">Zea mays</span>"
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        1 => array:4 [
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            3 => "RWC"
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    "highlights" => array:2 [
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      "resumen" => "<span id="abst0005" class="elsevierStyleSection elsevierViewall"><p id="spar0005" class="elsevierStyleSimplePara elsevierViewall"><ul class="elsevierStyleList" id="lis0005"><li class="elsevierStyleListItem" id="lsti0005"><span class="elsevierStyleLabel">&#8226;</span><p id="par0005" class="elsevierStylePara elsevierViewall"><span class="elsevierStyleItalic">A&#46; argentinense</span> Az19 mitigated maize growth reduction due to water deficit &#40;WD&#41; at V2&#46;</p></li><li class="elsevierStyleListItem" id="lsti0010"><span class="elsevierStyleLabel">&#8226;</span><p id="par0010" class="elsevierStylePara elsevierViewall">WD at flowering reduced maize yield less in Az19- than in Az39-inoculated plants&#46;</p></li><li class="elsevierStyleListItem" id="lsti0015"><span class="elsevierStyleLabel">&#8226;</span><p id="par0015" class="elsevierStylePara elsevierViewall">Az19 is a promising PGPR strain to be used as an inoculant in areas subjected to WD&#46;</p></li></ul></p></span>"
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        "resumen" => "<span id="abst0010" class="elsevierStyleSection elsevierViewall"><p id="spar0010" class="elsevierStyleSimplePara elsevierViewall">Water deficit constitutes a severe limitation to agricultural productivity&#46; In the context of sustainable crop production&#44; the potential of microbial biotechnology to increase plant drought tolerance and improve crop yields under adverse conditions is gaining relevance&#46; This work aimed to compare the performance of <span class="elsevierStyleItalic">Azospirillum</span><span class="elsevierStyleItalic">argentinense</span> strain Az19 to that of strain Az39&#44; the most widely used for commercial inoculants&#44; when inoculated in maize plants exposed to water deficit&#46; For this purpose&#44; greenhouse and field assays were conducted&#46; In the greenhouse experiment&#44; strain Az19 prevented the adverse effect of water deficit at V2 stage on maize growth&#46; Moreover&#44; the percentage of fertile plants and the ear weight decreased significantly under water deficits imposed at V2 and flowering in Az39-inoculated plants but not in Az19-inoculated plants&#46; In the first field trial with the commercial maize hybrid DOW DS 515 PW&#44; Az19-inoculated plants were those which better tolerated the water deficit imposed&#46; In the second field trial&#44; two maize genotypes with differential drought sensitivity &#40;LP 29<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 2542&#44; sensitive&#59; LP 882 &#40;923&#41;<span class="elsevierStyleHsp" style=""></span>&#215;<span class="elsevierStyleHsp" style=""></span>LP 4703&#44; tolerant&#41; were tested&#46; Higher tolerance to water deficit was detected in plants inoculated with <span class="elsevierStyleItalic">A&#46; argentinense</span> Az19&#44; with a noticeable effect on grain yield components in the sensitive genotype&#46; Based on these results&#44; we propose the use of <span class="elsevierStyleItalic">A&#46; argentinense</span> Az19 for the formulation of more targeted <span class="elsevierStyleItalic">Azospirillum</span>-based inoculants&#44; suitable for agroecological areas subjected to seasonal water deficits&#46;</p></span>"
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        "resumen" => "<span id="abst0015" class="elsevierStyleSection elsevierViewall"><p id="spar0015" class="elsevierStyleSimplePara elsevierViewall">El d&#233;ficit h&#237;drico constituye una severa limitaci&#243;n a la productividad agr&#237;cola&#46; En el marco de la producci&#243;n sostenible de cultivos&#44; la biotecnolog&#237;a microbiana est&#225; cobrando relevancia para aumentar la tolerancia a la sequ&#237;a y mejorar el rendimiento de los cultivos en condiciones adversas&#46; El prop&#243;sito de este trabajo fue comparar la acci&#243;n de la cepa de <span class="elsevierStyleItalic">Azospirillum argentinense</span> Az19&#44; con tolerancia in vitro a estresores abi&#243;ticos&#44; con la cepa Az39&#44; utilizada ampliamente para la formulaci&#243;n de inoculantes comerciales&#44; al inocularlas en plantas sometidas a d&#233;ficit h&#237;drico&#46; Se realizaron ensayos de invernadero y de campo&#46; En invernadero&#44; la cepa Az19 evit&#243; el impacto adverso del d&#233;ficit h&#237;drico en el estadio V2 sobre el crecimiento del ma&#237;z&#46; Adem&#225;s&#44; el porcentaje de plantas con espigas y el peso de la espiga disminuy&#243; significativamente con la restricci&#243;n h&#237;drica aplicada en V2 y en floraci&#243;n en plantas inoculadas con la cepa Az39&#44; pero no en las inoculadas con Az19&#46; En el primer ensayo de campo con el ma&#237;z h&#237;brido comercial DOW DS 515 PW las plantas inoculadas con Az19 fueron las que mejor toleraron la deficiencia h&#237;drica&#46; En el segundo ensayo de campo se utilizaron dos genotipos de ma&#237;z con sensibilidad diferencial a la sequ&#237;a&#46; La inoculaci&#243;n con Az19 condujo a una mayor tolerancia al d&#233;ficit h&#237;drico&#44; con un efecto detectable en algunos componentes del rendimiento en el genotipo sensible&#46; Sobre la base de estos resultados&#44; proponemos el empleo de <span class="elsevierStyleItalic">A&#46; argentinense</span> Az19 para la formulaci&#243;n de inoculantes basados en <span class="elsevierStyleItalic">Azospirillum</span> especialmente indicados para &#225;reas agroecol&#243;gicas que experimenten per&#237;odos de d&#233;ficit h&#237;drico&#46;</p></span>"
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          "en" => "<p id="spar0020" class="elsevierStyleSimplePara elsevierViewall">Effect of inoculation and water deficit at V2 on maize growth and physiological parameters &#40;greenhouse experiment&#41;&#46; Mean values and standard deviation determined at V3 are shown &#40;n<span class="elsevierStyleHsp" style=""></span>&#61;<span class="elsevierStyleHsp" style=""></span>12&#41;&#46; &#40;a&#41; Plant height&#59; &#40;b&#41; shoot fresh weight&#59; &#40;c&#41; shoot dry weight&#59; &#40;d&#41; relative water content of the last expanded leaf&#46; Control&#58; no water deficit&#59; WR&#58; water restriction&#59; NI&#58; non-inoculated&#46; Data was square root-transformed and analyzed by two-way ANOVA plus the Tukey&#39;s test&#46; Asterisks indicate significant differences between control and WR groups within each inoculation treatment &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#706;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p>"
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          "en" => "<p id="spar0025" class="elsevierStyleSimplePara elsevierViewall">Changes in maize height &#40;A&#41;&#44; percentage of empty ears &#40;B&#41;&#44; and yield &#40;C&#41; according to irrigation and inoculation treatments &#40;field assay 2&#41;&#46; Mean values and standard deviation determined at physiological maturity are shown&#46; Sampling was performed as described in &#8220;Materials and methods&#8221; section&#46; C&#58; full-irrigated control&#59; WR-Fl&#58; water restriction at flowering&#59; WR-Gf&#58; water restriction at grain filling&#59; GS&#58; drought-sensitive genotype&#59; GT&#58; drought-tolerant genotype&#46; Data was square root-transformed &#40;B&#41; or not transformed &#40;A and C&#41; and analyzed by two-way ANOVA plus the Tukey&#39;s test&#46; Different letters indicate significant differences between treatments irrespective of the plant genotype &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#706;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p>"
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          "leyenda" => "<p id="spar0035" class="elsevierStyleSimplePara elsevierViewall">Mean values obtained at physiological maturity are shown&#46; C&#58; irrigated on demand&#59; WR&#58; water restriction&#59; Ni&#58; not inoculated&#46; Asterisks indicate significant differences between C and WR groups within each inoculation treatment&#44; according to two-way ANOVA followed by &#352;&#237;d&#225;k&#39;s multiple range test &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#8804;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p>"
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                  \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Grain weight per plant &#40;g&#41;&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">C&#58;Ni&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">39&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>12&#46;34&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">10&#46;65<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>5&#46;07&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
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                  \t\t\t\t">5&#46;2<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>2&#46;12&nbsp;\t\t\t\t\t\t\n
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          "leyenda" => "<p id="spar0045" class="elsevierStyleSimplePara elsevierViewall">Mean values obtained at physiological maturity are shown&#46; Control&#58; full-irrigated&#59; WR-Fl&#58; water restriction at flowering&#59; WR-Gf&#58; water restriction at grain filling&#59; NI&#58; non-inoculated&#46; Different letters indicate significant differences between different water treatments within each inoculation group according to two-way ANOVA followed by Tukey&#39;s multiple range test &#40;<span class="elsevierStyleItalic">p</span><span class="elsevierStyleHsp" style=""></span>&#706;<span class="elsevierStyleHsp" style=""></span>0&#46;1&#41;&#46;</p>"
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                  \t\t\t\t" scope="col" style="border-bottom: 2px solid black">Water treatment&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">A&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">4&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1&#46;52&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n
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                  \t\t\t\t">4564<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1007&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t">B&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
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                  \t\t\t\t">29&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>13&#46;35&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
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                  \t\t\t\t">WR-Gf&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">B&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">Control&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">Az19&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t">10187<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>697&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t">A&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
                  \t\t\t\t\ttable-entry\n
                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t">2&#46;3<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1&#46;41&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
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                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n
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                  \t\t\t\t">WR-Fl&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
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                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t">5782<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>1009&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t  " align="left" valign="\n
                  \t\t\t\t\ttop\n
                  \t\t\t\t">B&nbsp;\t\t\t\t\t\t\n
                  \t\t\t\t</td><td class="td" title="\n
                  \t\t\t\t\ttable-entry\n
                  \t\t\t\t  " align="left" valign="\n
                  \t\t\t\t\ttop\n
                  \t\t\t\t">14&#46;9<span class="elsevierStyleHsp" style=""></span>&#177;<span class="elsevierStyleHsp" style=""></span>7&#46;09&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t ; entry_with_role_rowhead " align="left" valign="\n
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                  \t\t\t\t  " align="left" valign="\n
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                  \t\t\t\t">AB&nbsp;\t\t\t\t\t\t\n
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                  \t\t\t\t  " align="left" valign="\n
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                      "titulo" => "Response to saline stress and aquaporin expression in <span class="elsevierStyleItalic">Azospirillum</span>-inoculated barley seedlings"
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                            0 => "M&#46;S&#46; Zawoznik"
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                            2 => "M&#46;P&#46; Benavides"
                            3 => "S&#46; V&#225;zquez"
                            4 => "M&#46;D&#46; Groppa"
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                        "tituloSerie" => "Appl Microbiol Biotechnol"
                        "fecha" => "2011"
                        "volumen" => "90"
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        "identificador" => "xack692869"
        "titulo" => "Acknowledgments"
        "texto" => "<p id="par0210" class="elsevierStylePara elsevierViewall">Mar&#237;a Daniela Groppa and Guillermo Andr&#233;s Maroniche are researchers at the Consejo Nacional de Investigaciones Cient&#237;ficas y T&#233;cnicas &#40;CONICET&#41;&#46;</p><p id="par0215" class="elsevierStylePara elsevierViewall">This work has been funded by <span class="elsevierStyleGrantSponsor" id="gs1">Instituto Nacional de Tecnolog&#237;a Agropecuaria</span> &#40;<span class="elsevierStyleGrantNumber" refid="gs1">PNCYO-1127033</span>&#41; and the <span class="elsevierStyleGrantSponsor" id="gs2">University of Buenos Aires</span> &#40;<span class="elsevierStyleGrantNumber" refid="gs2">UBACYT 20020130200052BA</span>&#41;&#46;</p>"
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Article information

ISSN: 03257541
Original language: English

DOI:10.1016/j.ram.2023.01.002

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