RESUMO

Selenium (Se) is a beneficial element to higher plants. Application of Se at low concentrations enhances the antioxidant metabolism reducing the reactive oxygen species (ROS) generated by plant membrane cells. This study aimed to evaluate how the application of Se in the forms sodium selenate and sodium selenite regulates ROS scavenging in field-grown cowpea plants. Seven Se application rates (0; 2.5; 5; 10; 20; 40 and 60 g ha-1) of each of the two Se forms were applied to plants via the soil. Photosynthetic pigments concentration, gas exchange parameters, lipid peroxidation by malondialdehyde (MDA) concentration, hydrogen peroxide concentration, activity of catalase (CAT, EC:1.11.1.6), glutathione reductase (GR, EC:1.6.4.2), ascorbate peroxidase (APX, EC:1.11.1.11) and Se concentration in leaves and grains were evaluated. In general, Se application led to a decrease in chlorophyll a concentration whilst leading to an increase in chlorophyll b, indicating conservation of total chlorophyll concentration. Application of 2.5 g ha-1 of Se as selenate provided a notable increase in total chlorophyll and total carotenoids compared to the other application rates. Selenate and selenite application decreased lipid peroxidation. However, each Se source acted in a different pathway to combat ROS. While selenate showed more potential to increase activity of APX and GR, selenite showed a higher potential to increase CAT activity. The negative correlation between CAT and GR is indicative that both pathways might be activated under distinct circumstances. The more prominent activity of CAT under high rates of selenite resulted in a negative correlation of this enzyme with chlorophyll a and carotenoids. Both selenate and selenite application increased sucrose and total sugars concentration in leaves of cowpea plants. Overall, these results indicate that application of Se in cowpea under field conditions stimulates distinct pathways to scavenge ROS. This could prove beneficial to mitigate oxidative stress during plant development.

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Cowpea is an important source of protein, being the main subsistence crop in the northeast region of Brazil, where water restriction and salinity are limiting factors. Identification of saline stress tolerant varieties can aid crop expansion and improve crop yields. The objective was to study the tolerance of traditional varieties of cowpea to the salinity of irrigation water in the emergency and initial growth phase. The experiment was carried out in a greenhouse using a completely randomized design (DIC), in a 23 x 2 factorial scheme, composed of 23 cowpea varieties (Canapu Vermelho, Boquinha, Pingo de Ouro, Sempre Verde, Ceará, Baeta, Manteiga, Roxão, Costela de Vaca, Feijão Branco, Coruja, Rabo de Peba Branco, Sopinha, Canapu Branco, Lisão, Canapu Miúdo, Sempre Verde Ligeiro, Vagem Roxa, Ovo de Peru, Rabo de Peba Miúdo, Feijão Azul, Canário, and Paulistinha) and two levels of salinity of irrigation water (S1 = 0.5 and S2 = 5.5 dS m-¹), with four repetitions of 50 seeds. The seedlings were evaluated for emergence, vigor, salinity tolerance index, and dissimilarity. Increased salinity of irrigation water reduced emergence, vigor and dry matter accumulation of cowpea varieties. The cowpea varieties 'Lisão', 'Costela de Vaca', 'Canário', 'Feijão Branco', 'Ceará', and 'Boquinha' are the most tolerant to salinity, while the varieties 'Sempre Verde' and 'Manteiga' are the most sensitive to salinity in the emergency and initial growth phase.(AU)

O feijão-caupi é uma importante fonte de proteína, sendo a principal cultura de subsistência da região Nordeste do Brasil, onde a restrição de água e a salinidade são fatores limitantes. A identificação de variedades tolerantes ao estresse salino pode auxiliar na expansão da cultura, além de melhorar o rendimento dos cultivos. Objetivou-se estudar a tolerância de variedades crioulas de feijão-caupi à salinidade da água de irrigação na fase de emergência e crescimento inicial. O experimento foi realizado em casa de vegetação, usando o delineamento inteiramente casualizado (DIC), em esquema fatorial 23x 2, composto de 23 variedades de feijão-caupi (Canapu Vermelho, Boquinha, Pingo de Ouro, Sempre Verde, Ceará, Baeta, Manteiga, Roxão, Costela de Vaca, Feijão Branco, Coruja, Rabo de Peba Branco, Sopinha, Canapu Branco, Lisão, Canapu Miúdo, Sempre Verde Ligeiro, Vagem Roxa, Ovo de Peru, Rabo de Peba Miúdo, Feijão Azul, Canário e Paulistinha), e dois níveis de salinidade da água de irrigação (S1= 0,5 e S2 = 5,5 dS m-1), com quatro repetições de 50 sementes. As plântulas foram avaliadas quanto à emergência, o vigor, o índice de tolerância à salinidade e à dissimilaridade. O aumento da salinidade da água de irrigação reduziu a emergência, o vigor e acúmulo de massa seca das variedades de feijão-caupi. As variedades de feijão-caupi Lisão, Costela de Vaca, Canário, Feijão Branco, Ceará e Boquinha são as mais tolerantes à salinidade, enquanto que as variedades Sempre Verde e Manteiga são as mais sensíveis à salinidade na fase de emergência e crescimento inicial.(AU)

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RESUMO

Cowpea is an important source of protein, being the main subsistence crop in the northeast region of Brazil, where water restriction and salinity are limiting factors. Identification of saline stress tolerant varieties can aid crop expansion and improve crop yields. The objective was to study the tolerance of traditional varieties of cowpea to the salinity of irrigation water in the emergency and initial growth phase. The experiment was carried out in a greenhouse using a completely randomized design (DIC), in a 23 x 2 factorial scheme, composed of 23 cowpea varieties (Canapu Vermelho, Boquinha, Pingo de Ouro, Sempre Verde, Ceará, Baeta, Manteiga, Roxão, Costela de Vaca, Feijão Branco, Coruja, Rabo de Peba Branco, Sopinha, Canapu Branco, Lisão, Canapu Miúdo, Sempre Verde Ligeiro, Vagem Roxa, Ovo de Peru, Rabo de Peba Miúdo, Feijão Azul, Canário, and Paulistinha) and two levels of salinity of irrigation water (S1 = 0.5 and S2 = 5.5 dS m-¹), with four repetitions of 50 seeds. The seedlings were evaluated for emergence, vigor, salinity tolerance index, and dissimilarity. Increased salinity of irrigation water reduced emergence, vigor and dry matter accumulation of cowpea varieties. The cowpea varieties 'Lisão', 'Costela de Vaca', 'Canário', 'Feijão Branco', 'Ceará', and 'Boquinha' are the most tolerant to salinity, while the varieties 'Sempre Verde' and 'Manteiga' are the most sensitive to salinity in the emergency and initial growth phase.

O feijão-caupi é uma importante fonte de proteína, sendo a principal cultura de subsistência da região Nordeste do Brasil, onde a restrição de água e a salinidade são fatores limitantes. A identificação de variedades tolerantes ao estresse salino pode auxiliar na expansão da cultura, além de melhorar o rendimento dos cultivos. Objetivou-se estudar a tolerância de variedades crioulas de feijão-caupi à salinidade da água de irrigação na fase de emergência e crescimento inicial. O experimento foi realizado em casa de vegetação, usando o delineamento inteiramente casualizado (DIC), em esquema fatorial 23x 2, composto de 23 variedades de feijão-caupi (Canapu Vermelho, Boquinha, Pingo de Ouro, Sempre Verde, Ceará, Baeta, Manteiga, Roxão, Costela de Vaca, Feijão Branco, Coruja, Rabo de Peba Branco, Sopinha, Canapu Branco, Lisão, Canapu Miúdo, Sempre Verde Ligeiro, Vagem Roxa, Ovo de Peru, Rabo de Peba Miúdo, Feijão Azul, Canário e Paulistinha), e dois níveis de salinidade da água de irrigação (S1= 0,5 e S2 = 5,5 dS m-1), com quatro repetições de 50 sementes. As plântulas foram avaliadas quanto à emergência, o vigor, o índice de tolerância à salinidade e à dissimilaridade. O aumento da salinidade da água de irrigação reduziu a emergência, o vigor e acúmulo de massa seca das variedades de feijão-caupi. As variedades de feijão-caupi Lisão, Costela de Vaca, Canário, Feijão Branco, Ceará e Boquinha são as mais tolerantes à salinidade, enquanto que as variedades Sempre Verde e Manteiga são as mais sensíveis à salinidade na fase de emergência e crescimento inicial.

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O presente trabalho objetivou analisar o desempenho fisiológico de sementes de feijão-caupi cv. BRS Pajeú submetidas à diferentes potenciais e agentes osmóticos, permitindo o conhecimento sobre como o estresse influencia na germinação e vigor destas sementes. O experimento foi conduzido no delineamento inteiramente casualizado, com os tratamentos em arranjo fatorial sendo dois agentes osmóticos e cinco potenciais osmóticos (2 × 5), com quatro repetições. Os resultados obtidos foram submetidos à análise de variância, e quando constatados efeitos significativos, foram submetidos à análise de regressão. As condições de estresse salino prejudicaram a germinação e o vigor das sementes de feijão-caupi cv. BRS Pajeú. A diminuição do potencial osmótico das soluções, reduz o desenvolvimento e o ganho de massa fresca e seca das plântulas de feijão-caupi cv. BRS Pajeú, independente da substância utilizada para indução do estresse. O cloreto de sódio (NaCl) promoveu efeitos mais drásticos sobre o crescimento das plântulas, quando comparado ao cloreto de potássio (KCl). As sementes de feijão-caupi cv BRS Pajeú submetidas à solução de NaCl com potenciais osmóticos inferiores a -0,35 MPa, apresentou porcentagem inferior a 80% de germinação.(AU)

The aim of this work was to evaluate the germination physiological of cowpea seed cv. BRS Pajeú submitted to different potentials and osmotic agents, allowing the knowledge about how the stress effects the germination and vigor of these seeds. The experiment was conducted in a completely randomized design, with treatments in five different potentials and two osmotic agents (2 ×5) factorial arrangement, with four replications. The results were submitted to variance analysis, and when significant effects were verified, they were submitted to regression analysis. The saline stress conditions impaired germination and vigor of cowpea seeds. The decrease in the osmotic potential of the solutions reduces the development and fresh and dry mass gain of cowpea cv. BRS Pajeú, independent of the substance used for stress induction. Sodium chloride (NaCl) promoted more drastic effects on seedling growth when compared to potassium chloride (KCl). The cowpea seeds submitted to NaCl solution with potentials inferior to -0.35 MPa, presented less than 80% of germination.(AU)

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O presente trabalho objetivou analisar o desempenho fisiológico de sementes de feijão-caupi cv. BRS Pajeú submetidas à diferentes potenciais e agentes osmóticos, permitindo o conhecimento sobre como o estresse influencia na germinação e vigor destas sementes. O experimento foi conduzido no delineamento inteiramente casualizado, com os tratamentos em arranjo fatorial sendo dois agentes osmóticos e cinco potenciais osmóticos (2 × 5), com quatro repetições. Os resultados obtidos foram submetidos à análise de variância, e quando constatados efeitos significativos, foram submetidos à análise de regressão. As condições de estresse salino prejudicaram a germinação e o vigor das sementes de feijão-caupi cv. BRS Pajeú. A diminuição do potencial osmótico das soluções, reduz o desenvolvimento e o ganho de massa fresca e seca das plântulas de feijão-caupi cv. BRS Pajeú, independente da substância utilizada para indução do estresse. O cloreto de sódio (NaCl) promoveu efeitos mais drásticos sobre o crescimento das plântulas, quando comparado ao cloreto de potássio (KCl). As sementes de feijão-caupi cv BRS Pajeú submetidas à solução de NaCl com potenciais osmóticos inferiores a -0,35 MPa, apresentou porcentagem inferior a 80% de germinação.

The aim of this work was to evaluate the germination physiological of cowpea seed cv. BRS Pajeú submitted to different potentials and osmotic agents, allowing the knowledge about how the stress effects the germination and vigor of these seeds. The experiment was conducted in a completely randomized design, with treatments in five different potentials and two osmotic agents (2 ×5) factorial arrangement, with four replications. The results were submitted to variance analysis, and when significant effects were verified, they were submitted to regression analysis. The saline stress conditions impaired germination and vigor of cowpea seeds. The decrease in the osmotic potential of the solutions reduces the development and fresh and dry mass gain of cowpea cv. BRS Pajeú, independent of the substance used for stress induction. Sodium chloride (NaCl) promoted more drastic effects on seedling growth when compared to potassium chloride (KCl). The cowpea seeds submitted to NaCl solution with potentials inferior to -0.35 MPa, presented less than 80% of germination.

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Correlations between the transcriptional responses of genes that encode superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), and peroxiredoxin (Prx) enzymes and Colletotrichum gloeosporioides development in cowpea leaves were assessed. Each of these genes is involved in the redox metabolism and hydrogen peroxide balance. Although electron microscopy revealed that conidia adhered to and germinated on the leaf cuticle, the inoculated cowpea leaves did not show any characteristic anthracnose symptoms. The adhered and germinated conidia showed irregular surfaces and did not develop further. This was apparently due to increased leaf H2O2 levels in response to inoculation with C. gloeosporioides. During the early stages post inoculation, cowpea leaves elevated the H2O2 content and modulated the defense gene expression, as well as associated pathways. During the later stages, the increased expression of the CuZnSODI and CuZnSODII genes suggested an active superoxide dismutation to further elevate H2O2 levels, which indicated that higher H2O2 content may function as a toxic agent that kills the fungus. The second increase in H2O2 production above the threshold level was correlated with the expression of the APXI, CATI, CATII, PrxIIBCD, and PrxIIE genes, which resulted in a coordinated pattern to establish an appropriate balance between H2O2 generation and scavenging. Therefore, appropriate H2O2 content in cowpea leaves inhibited C. gloeosporioides development and maintained intracellular redox homeostasis to avoid uncontrolled programmed cell death and necrosis in cowpea leaves.

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ABSTRACT The leguminous inoculation with nodule-inducing bacteria that perform biological nitrogen fixation is a good example of an "eco-friendly agricultural practice". Bradyrhizobium strains BR 3267 and BR 3262 are recommended for cowpea (Vigna unguiculata) inoculation in Brazil and showed remarkable responses; nevertheless neither strain was characterized at species level, which is our goal in the present work using a polyphasic approach. The strains presented the typical phenotype of Bradyrhizobium with a slow growth and a white colony on yeast extract-mannitol medium. Strain BR 3267 was more versatile in its use of carbon sources compared to BR 3262. The fatty acid composition of BR 3267 was similar to the type strain of Bradyrhizobium yuanmingense; while BR 3262 was similar to Bradyrhizobium elkanii and Bradyrhizobium pachyrhizi. Phylogenetic analyses based on 16S rRNA and three housekeeping genes placed both strains within the genus Bradyrhizobium: strain BR 3267 was closest to B. yuanmingense and BR 3262 to B. pachyrhizi. Genome average nucleotide identity and DNA-DNA reassociation confirmed the genomic identification of B. yuanmingense BR 3267 and B. pachyrhizi BR 3262. The nodC and nifH gene analyses showed that strains BR 3267 and BR 3262 hold divergent symbiotic genes. In summary, the results indicate that cowpea can establish effective symbiosis with divergent bradyrhizobia isolated from Brazilian soils.

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RESUMO

The leguminous inoculation with nodule-inducing bacteria that perform biological nitrogen fixation is a good example of an "eco-friendly agricultural practice". Bradyrhizobium strains BR 3267 and BR 3262 are recommended for cowpea (Vigna unguiculata) inoculation in Brazil and showed remarkable responses; nevertheless neither strain was characterized at species level, which is our goal in the present work using a polyphasic approach. The strains presented the typical phenotype of Bradyrhizobium with a slow growth and a white colony on yeast extract-mannitol medium. Strain BR 3267 was more versatile in its use of carbon sources compared to BR 3262. The fatty acid composition of BR 3267 was similar to the type strain of Bradyrhizobium yuanmingense; while BR 3262 was similar to Bradyrhizobium elkanii and Bradyrhizobium pachyrhizi. Phylogenetic analyses based on 16S rRNA and three housekeeping genes placed both strains within the genus Bradyrhizobium: strain BR 3267 was closest to B. yuanmingense and BR 3262 to B. pachyrhizi. Genome average nucleotide identity and DNA-DNA reassociation confirmed the genomic identification of B. yuanmingense BR 3267 and B. pachyrhizi BR 3262. The nodC and nifH gene analyses showed that strains BR 3267 and BR 3262 hold divergent symbiotic genes. In summary, the results indicate that cowpea can establish effective symbiosis with divergent bradyrhizobia isolated from Brazilian soils.

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KEY MESSAGE: The seed treatment of a CPSMV-susceptible cowpea genotype with the mutagenic agent EMS generated mutagenized resistant plantlets that respond to the virus challenge by activating biochemical and physiological defense mechanisms. Cowpea is an important crop that makes major nutritional contributions particularly to the diet of the poor population worldwide. However, its production is low, because cowpea is naturally exposed to several abiotic and biotic stresses, including viral agents. Cowpea severe mosaic virus (CPSMV) drastically affects cowpea grain production. This study was conducted to compare photosynthetic and biochemical parameters of a CPSMV-susceptible cowpea (CE-31 genotype) and its derived ethyl methanesulfonate-mutagenized resistant plantlets, both challenged with CPSMV, to shed light on the mechanisms of virus resistance. CPSMV inoculation was done in the fully expanded secondary leaves, 15 days after planting. At 7 days post-inoculation, in vivo photosynthetic parameters were measured and leaves collected for biochemical analysis. CPSMV-inoculated mutagenized-resistant cowpea plantlets (MCPI) maintained higher photosynthesis index, chlorophyll, and carotenoid contents in relation to the susceptible (CE-31) CPSMV-inoculated cowpea (CPI). Visually, the MCPI leaves did not exhibit any viral symptoms neither the presence of the virus as examined by RT-PCR. In addition, MCPI showed higher SOD, GPOX, chitinase, and phenylalanine ammonia lyase activities, H2O2, phenolic contents, and cell wall lignifications, but lower CAT and APX activities in comparison to CPI. All together, these photosynthetic and biochemical changes might have contributed for the CPSMS resistance of MCPI. Contrarily, CPI plantlets showed CPSMV accumulation, severe disease symptoms, reduction in the photosynthesis-related parameters, chlorophyll, carotenoid, phenolic compound, and H2O2 contents, in addition to increased ß-1,3-glucanase, and catalase activities that might have favored viral infection.

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The strain BR 3267 is a nitrogen-fixing symbiotic bacteria isolated from soil of semi-arid area of Brazilian Northeast using cowpea as the trap plant. This strain is used as commercial inoculant for cowpea and presents high efficient in nitrogen fixation as consequence of its adaptation potential to semi-arid conditions. We report here the draft genome sequence of Bradyrhizobium sp. strain BR 3267, an elite bacterium used as inoculant for cowpea. Whole genome sequencing of BR 3267 using Illumina MiSeq sequencing technology has 55 scaffolds with a total genome size of 7,904,309bp and C+G 63%. Annotation was added by the RAST prokaryotic genome annotation service and has shown 7314 coding sequences and 52 RNA genes.

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The strain BR 3262 was isolated from nodule of cowpea (Vigna unguiculata L. Walp) growing in soil of the Atlantic Forest area in Brazil and it is reported as an efficient nitrogen fixing bacterium associated to cowpea. Firstly, this strain was assigned as Bradyrhizobium elkanii, however, recently a more detailed genetic and molecular characterization has indicated it could be a Bradyrhizobium pachyrhizi species. We report here the draft genome sequence of B. pachyrhizi strain BR 3262, an elite bacterium used as inoculant for cowpea. The whole genome with 116 scaffolds, 8,965,178bp and 63.8% of C+G content for BR 3262 was obtained using Illumina MiSeq sequencing technology. Annotation was added by the RAST prokaryotic genome annotation service and shown 8369 coding sequences, 52 RNAs genes, classified in 504 subsystems.

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This study aimed to verify that a Bayesian approach could be used for the selection of upright cowpea genotypes with high adaptability and phenotypic stability, and the study also evaluated the efficiency of using informative and minimally informative a priori distributions. Six trials were conducted in randomized blocks, and the grain yield of 17 upright cowpea genotypes was assessed. To represent the minimally informative a priori distributions, a probability distribution with high variance was used, and a meta-analysis concept was adopted to represent the informative a priori distributions. Bayes factors were used to conduct comparisons between the a priori distributions. The Bayesian approach was effective for selection of upright cowpea genotypes with high adaptability and phenotypic stability using the Eberhart and Russell method. Bayes factors indicated that the use of informative a priori distributions provided more accurate results than minimally informative a priori distributions.

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O presente trabalho teve como objetivo avaliar o efeito alelopático do óleo essencial de plantas de carqueja, Baccharis trimera (Less.) DC., sobre a germinação de sementes de feijão Vigna unguiculata (L.) Walp. Foi avaliado o efeito do óleo essencial de B. trimera sobre V. unguiculata nas dosagens 20 μL, 15 μL, 10 μL, 5 μL e testemunha. A qualidade fisiológica das sementes foi determinada pela porcentagem de emergência, velocidade de emergência e índice de velocidade de emergência. O delineamento experimental foi o inteiramente casualizado, em esquema fatorial 5 x 2 com cinco repetições. Não foi observado efeito inibidor do óleo essencial de B. trimera na germinação de sementes de feijão caupi, caracterizando-se como de efeito alelopático benéfico. De acordo com os resultados obtidos, o óleo essencial de B. trimera revelou-se eficiente na manutenção da viabilidade dessas sementes.

This research aimed to evaluate the allellopathic effect of essential oils of the Baccharis trimera (Less.) DC. plants on the seeds germination of cowpea Vigna unguiculata (L.) Walp. The effect of B. trimera essential oil on V. unguiculata was evaluated at levels of 20 μL, 15 μL, 10 μL, 5 μL and control. The physiologic quality of the seeds was determined by percentage emergence, the rate of speed emergence and speed emergence index. The data analysis was carried out using an entirely randomized design, in a 5 x 2 factorial scheme with five repetitions. An inhibitory effect of B. trimera essential oil on bean seeds germination was not observed, which is characterized as a beneficial allelopathic effect. Based on the results, the B. trimera essential oil proved efficient in the viability maintenance of these seeds.

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A adubação fosfatada e as formas de aplicação de fertilizantes fosfatados são fatores importantes no manejo das culturas. O objetivo do presente trabalho foi avaliar os efeitos de doses de fósforo e formas de aplicação de fertilizante fosfatado na cultura do feijão-caupi (Vigna unguiculata (L.) Walp) em solo de cerrado do Estado de Roraima. O experimento foi realizado em condições de campo, em Latossolo Amarelo distrocoeso, no Campus do Cauamé da Universidade Federal de Roraima. O delineamento experimental foi o de parcelas subdivididas, com as parcelas principais organizadas em blocos casualizados, com três repetições. As parcelas foram constituídas pelas formas de aplicação do fertilizante fosfatado: a) Lanço - L; b) Sulco simples - SS e c) Sulco duplo - SD. As subparcelas corresponderam às doses da adubação fosfatada: 0; 0,5; 1,0; 1,5 e 2,0 vezes a dose recomendada para adubação de manutenção do feijão-caupi. Houve efeito das doses de fósforo sobre o crescimento e absorção de fósforo pelas plantas. O maior crescimento, produção de vagem por planta, de grãos e absorção de fósforo ocorreu com a aplicação do fertilizante fosfatado aplicado em sulcos duplos.

Phosphorus fertilization and ways of phosphate fertilizers application are important factors considering crop management. This study aimed at assessing the effect of phosphorus rates and phosphate fertilizers application forms on growth and phosphorus uptake of cowpea plants cropped on cerrado soil in the State of Roraima, Brazil. An experiment took place under field conditions, on a Yellow Latosol, at Cauamé Campus of Roraima's Federal University, located in the municipality of Boa Vista/RR. The experimental design was split-plot, in a completely randomized blocks, with three replicates. The plots consisted of the following phosphoric fertilizers application forms: a) broadcasting; b) single furrow and c) double furrow. Sub-plots corresponded to phosphorus doses: 0, 0,5; 1,0; 1,5, and 2,0 times the recommended dose for cowpea maintenance fertilization (60 kg ha-1 of P2O5). There was a significant effect of P rates over plant growth and plant uptake. Application of fertilizer in double furrows promoted higher plant growth, grain yield, and plant phosphorous uptake.

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