RESUMO
MAIN CONCLUSION: Inbred line 11-133 of popcorn showed the lowest apoplast Al and total Al concentrations and Al-lumogallion complex, associated with a more efficient antioxidant system, mainly due to glutathione metabolism. Popcorn (Zea mays L. var. everta) is largely intended for human consumption. About 40% of the world's arable soils are acidic. In soils acidic, aluminum (Al) ionizes producing the trivalent cation, which is highly toxic to plants. Hence, this work aimed to: (1) evaluate the Al toxicity sites and its effect on the structure of the root tips, (2) quantify Al concentrations in the apoplast and symplast of the roots, and (3) to elucidate the modulation on the activity of antioxidant enzymes and metabolites of the ascorbate-glutathione cycle in two popcorn inbred lines (ILs) 11-133 and 11-60, classified as tolerant and sensitive to this metal, respectively. Aluminum toxicity did not affect the shoot growth; however, there was a yellowing of the oldest leaf blade only in 11-60. The better performance of 11-133 is related to lower apoplastic and total Al concentrations and Al accumulation in the root associated with a lower fluorescence of Al-lumogallion complex at the root tip, indicating the presence of mechanisms of chelation with this metal. Consequently, this IL showed less change in root morphoanatomy and lower reactive oxygen species and malondialdehyde content, which are associated with a more efficient enzymatic and non-enzymatic system, mainly due to the higher content of the glutathione metabolite and the higher activities of superoxide dismutase, monodehydroascorbate reductase, dehydroascorbate reductase, γ-glutamylcysteine synthetase, and glutathione peroxidase enzymes. Thus, these findings illustrated above indicate how internal mechanisms of detoxification respond to Al in popcorn, which can be used as tolerance biomarkers.
Assuntos
Alumínio , Antioxidantes , Humanos , Antioxidantes/metabolismo , Alumínio/toxicidade , Estresse Oxidativo , Catalase/metabolismo , Ácido Ascórbico/metabolismo , Oxirredução , Glutationa , Solo , Raízes de Plantas/metabolismoRESUMO
Functional enrichment analysis is a cornerstone in bioinformatics as it makes possible to identify functional information by using a gene list as source. Different tools are available to compare gene ontology (GO) terms, based on a directed acyclic graph structure or content-based algorithms which are time-consuming and require a priori information of GO terms. Nevertheless, quantitative procedures to compare GO terms among gene lists and species are not available. Here we present a computational procedure, implemented in R, to infer functional information derived from comparative strategies. GOCompare provides a framework for functional comparative genomics starting from comparable lists from GO terms. The program uses functional enrichment analysis (FEA) results and implement graph theory to identify statistically relevant GO terms for both, GO categories and analyzed species. Thus, GOCompare allows finding new functional information complementing current FEA approaches and extending their use to a comparative perspective. To test our approach GO terms were obtained for a list of aluminum tolerance-associated genes in Oryza sativa subsp. japonica and their orthologues in Arabidopsis thaliana. GOCompare was able to detect functional similarities for reactive oxygen species and ion binding capabilities which are common in plants as molecular mechanisms to tolerate aluminum toxicity. Consequently, the R package exhibited a good performance when implemented in complex datasets, allowing to establish hypothesis that might explain a biological process from a functional perspective, and narrowing down the possible landscapes to design wet lab experiments.
Assuntos
Alumínio , Arabidopsis , Genômica/métodos , Biologia Computacional/métodos , Algoritmos , Ontologia Genética , Arabidopsis/genéticaRESUMO
Crop tolerance to multiple abiotic stresses has long been pursued as a Holy Grail in plant breeding efforts that target crop adaptation to tropical soils. On tropical, acidic soils, aluminum (Al) toxicity, low phosphorus (P) availability and drought stress are the major limitations to yield stability. Molecular breeding based on a small suite of pleiotropic genes, particularly those with moderate to major phenotypic effects, could help circumvent the need for complex breeding designs and large population sizes aimed at selecting transgressive progeny accumulating favorable alleles controlling polygenic traits. The underlying question is twofold: do common tolerance mechanisms to Al toxicity, P deficiency and drought exist? And if they do, will they be useful in a plant breeding program that targets stress-prone environments. The selective environments in tropical regions are such that multiple, co-existing regulatory networks may drive the fixation of either distinctly different or a smaller number of pleiotropic abiotic stress tolerance genes. Recent studies suggest that genes contributing to crop adaptation to acidic soils, such as the major Arabidopsis Al tolerance protein, AtALMT1, which encodes an aluminum-activated root malate transporter, may influence both Al tolerance and P acquisition via changes in root system morphology and architecture. However, trans-acting elements such as transcription factors (TFs) may be the best option for pleiotropic control of multiple abiotic stress genes, due to their small and often multiple binding sequences in the genome. One such example is the C2H2-type zinc finger, AtSTOP1, which is a transcriptional regulator of a number of Arabidopsis Al tolerance genes, including AtMATE and AtALMT1, and has been shown to activate AtALMT1, not only in response to Al but also low soil P. The large WRKY family of transcription factors are also known to affect a broad spectrum of phenotypes, some of which are related to acidic soil abiotic stress responses. Hence, we focus here on signaling proteins such as TFs and protein kinases to identify, from the literature, evidence for unifying regulatory networks controlling Al tolerance, P efficiency and, also possibly drought tolerance. Particular emphasis will be given to modification of root system morphology and architecture, which could be an important physiological "hub" leading to crop adaptation to multiple soil-based abiotic stress factors.
RESUMO
Root growth is reduced in soils with low pH [H+] and abundant soluble aluminum [Al3+], which can be a consequence of the interaction between Al3+ and cell wall composition. The competition between Al3+ and Ca2+ toward binding to pectin molecules was evaluated in roots of Urochloa decumbens, an African grass highly adapted to acidic Al-rich soils. Variations in the composition and distribution of pectins can change the extensibility, rigidity, porosity, and adhesive properties of plant cell walls, which were tested in seedlings of U. decumbens exposed to pH 3.5, 4.5 and 5.8 and to 0, 80, 160 and 320 µM of Al3+ for 80h. Root growth corroborated that U. decumbens is very tolerant to soil acidity, with effective reduction of root growth only at pH 3.5. Immunocytochemical approaches demonstrated variations in pectin composition induced both by Al3+ and by H+ in root tissues and zones. Based on the usual linkage between Ca2+ and pectins, Density Functional Theory (DFT) analyses indicated that Al3+ bound easier to pectins than Ca2+ did, leading to the formation of more Al3+-pectate complexes than Ca2+-pectate complexes, which resulted in higher rigidity of cell walls, and hampered cell extension.
Assuntos
Alumínio/metabolismo , Cálcio/metabolismo , Pectinas/metabolismo , Raízes de Plantas/metabolismo , Poaceae/metabolismo , Parede Celular , Teoria da Densidade Funcional , Imuno-HistoquímicaRESUMO
High concentrations of heavy metal (HM) ions impact agronomic staple crop production in acid soils (pH ≤ 5) due to their cytotoxic, genotoxic, and mutagenic effects. Among cytotoxic ions, the trivalent aluminum cation (Al3+) formed by solubilization of aluminum (Al) into acid soils, is one of the most abundant and toxic elements under acidic conditions. In recent years, several studies have elucidated the different signal transduction pathways involved in HM responses, identifying complementary genetic mechanisms conferring tolerance to plants. Although epigenetics has become more relevant in abiotic stress studies, epigenetic mechanisms underlying plant responses to HM stress remain poorly understood. This review describes the main epigenetic mechanisms related to crop responses during stress conditions, specifically, the molecular evidence showing how epigenetics is at the core of plant adaptation responses to HM ions. We highlight the epigenetic mechanisms that induce Al tolerance. Likewise, we analyze the pivotal relationship between epigenetic and genetic factors associated with HM tolerance. Finally, using rice as a study case, we performed a general analysis over previously whole-genome bisulfite-seq published data. Specific genes related to Al tolerance, measured in contrasting tolerant and susceptible rice varieties, exhibited differences in DNA methylation frequency. The differential methylation patterns could be associated with epigenetic regulation of rice responses to Al stress, highlighting the major role of epigenetics over specific abiotic stress responses.
RESUMO
Presence of the ALMT1 (aluminum-activated malate transporter) gene confers resistance to aluminum toxicity in Triticum aestivum (common wheat). No resistant cultivars of Triticum turgidum ssp. Durum Desf. (durum wheat) have been registered in Brazil. The aim of this study was to map the ALMT1 through application of the FISH (fluorescence in situ hybridization) technique in five wheat genotypes, common and durum, from the Active Germplasm Bank (AGB) of the Instituto Agronômico (IAC): BH 1146, P19, P33, Anahuac, and IAC 1003. FISH-ALMT1 signals were registered in Anahuac (sensitive) chromosomes and in BH 1146, P19, and P33 (resistant) chromosomes. In the three resistant genotypes, a characteristic double FISH signal was found, located in different chromosomes of the complements: in BH 1146 in chromosome 5D, in P19 in 3B, and in P33 in 6B. This FISH - ALMT1 mapping allows for introgression of aluminum resistance in sensitive cultivars through breeding programs using introgression lines containing these carrier chromosomes.(AU)
Assuntos
Triticum/genética , Análise Citogenética/métodos , Alumínio/toxicidade , Hibridização in Situ FluorescenteRESUMO
Presence of the ALMT1 (aluminum-activated malate transporter) gene confers resistance to aluminum toxicity in Triticum aestivum (common wheat). No resistant cultivars of Triticum turgidum ssp. Durum Desf. (durum wheat) have been registered in Brazil. The aim of this study was to map the ALMT1 through application of the FISH (fluorescence in situ hybridization) technique in five wheat genotypes, common and durum, from the Active Germplasm Bank (AGB) of the Instituto Agronômico (IAC): BH 1146, P19, P33, Anahuac, and IAC 1003. FISH-ALMT1 signals were registered in Anahuac (sensitive) chromosomes and in BH 1146, P19, and P33 (resistant) chromosomes. In the three resistant genotypes, a characteristic double FISH signal was found, located in different chromosomes of the complements: in BH 1146 in chromosome 5D, in P19 in 3B, and in P33 in 6B. This FISH - ALMT1 mapping allows for introgression of aluminum resistance in sensitive cultivars through breeding programs using introgression lines containing these carrier chromosomes.
Assuntos
Alumínio/toxicidade , Análise Citogenética/métodos , Triticum/genética , Hibridização in Situ FluorescenteRESUMO
Aluminum (Al) toxicity on acidic soils significantly damages plant roots and inhibits root growth. Hence, crops intoxicated by Al become more sensitive to drought stress and mineral nutrient deficiencies, particularly phosphorus (P) deficiency, which is highly unavailable on tropical soils. Advances in our understanding of the physiological and genetic mechanisms that govern plant Al resistance have led to the identification of Al resistance genes, both in model systems and in crop species. It has long been known that Al resistance has a beneficial effect on crop adaptation to acidic soils. This positive effect happens because the root systems of Al resistant plants show better development in the presence of soil ionic Al3+ and are, consequently, more efficient in absorbing sub-soil water and mineral nutrients. This effect of Al resistance on crop production, by itself, warrants intensified efforts to develop and implement, on a breeding scale, modern selection strategies to profit from the knowledge of the molecular determinants of plant Al resistance. Recent studies now suggest that Al resistance can exert pleiotropic effects on P acquisition, potentially expanding the role of Al resistance on crop adaptation to acidic soils. This appears to occur via both organic acid (OA)- and non-OA transporters governing a joint, iron-dependent interplay between Al resistance and enhanced P uptake, via changes in root system architecture. Current research suggests this interplay to be part of a P stress response, suggesting that this mechanism could have evolved in crop species to improve adaptation to acidic soils. Should this pleiotropism prove functional in crop species grown on acidic soils, molecular breeding based on Al resistance genes may have a much broader impact on crop performance than previously anticipated. To explore this possibility, here we review the components of this putative effect of Al resistance genes on P stress responses and P nutrition to provide the foundation necessary to discuss the recent evidence suggesting pleiotropy as a genetic linkage between Al resistance and P efficiency. We conclude by exploring what may be needed to enhance the utilization of Al resistance genes to improve crop production on acidic soils.
RESUMO
In acid soils, toxic aluminum ions inhibit plant root growth. In order to discriminate aluminum (Al) tolerance, trustful screening techniques are required. In this study, 20 wheat cultivars, showing different levels of Al tolerance, were evaluated in a short-term soil experiment to access their relative root length (RRL). Moreover, the alleles of two important genes (TaALMT1 and TaMATE1B) for Al tolerance in wheat were discriminated. Both of these genes encode membrane transporters responsible for the efflux of organic acids by the root apices that are thought to confer tolerance by chelating Al. Genotypes showing TaALMT1 alleles V and VI and an insertion at the TaMATE1B promoter were among the ones showing greater RRL. Mechanisms of Al tolerance, which are not associated with organic acid efflux, can be potentially present in two cultivars showing greater RRL among the ones carrying inferior TaALMT1 and TaMATE1B alleles. The RRL data were highly correlated with wheat performance in acid soil at three developmental stages, tillering (r = 0.93, p 0.001), silking (r = 0.91, p 0.001) and maturation (r = 0.90, p 0.001), as well as with the classification index of aluminum toxicity in the field (r = 0.92, p 0.001). Since the RRL was obtained after only six days of growth and it is highly correlated with plant performance in acid soil under field conditions, the short-term experiment detailed here is an efficient and rapid method for reliable screening of wheat Al tolerance.(AU)
Assuntos
Triticum/química , Acidez do Solo/análise , Raízes de Plantas/crescimento & desenvolvimento , Alumínio , Malatos , CitratosRESUMO
In acid soils, toxic aluminum ions inhibit plant root growth. In order to discriminate aluminum (Al) tolerance, trustful screening techniques are required. In this study, 20 wheat cultivars, showing different levels of Al tolerance, were evaluated in a short-term soil experiment to access their relative root length (RRL). Moreover, the alleles of two important genes (TaALMT1 and TaMATE1B) for Al tolerance in wheat were discriminated. Both of these genes encode membrane transporters responsible for the efflux of organic acids by the root apices that are thought to confer tolerance by chelating Al. Genotypes showing TaALMT1 alleles V and VI and an insertion at the TaMATE1B promoter were among the ones showing greater RRL. Mechanisms of Al tolerance, which are not associated with organic acid efflux, can be potentially present in two cultivars showing greater RRL among the ones carrying inferior TaALMT1 and TaMATE1B alleles. The RRL data were highly correlated with wheat performance in acid soil at three developmental stages, tillering (r = 0.93, p 0.001), silking (r = 0.91, p 0.001) and maturation (r = 0.90, p 0.001), as well as with the classification index of aluminum toxicity in the field (r = 0.92, p 0.001). Since the RRL was obtained after only six days of growth and it is highly correlated with plant performance in acid soil under field conditions, the short-term experiment detailed here is an efficient and rapid method for reliable screening of wheat Al tolerance.
Assuntos
Acidez do Solo/análise , Raízes de Plantas/crescimento & desenvolvimento , Triticum/química , Alumínio , Citratos , MalatosRESUMO
Un problema para la producción agrícola en la Orinoquia colombiana, son los 4,5 millones de hectáreas con altos contenidos de aluminio. Genotipos de diferentes especies presentan niveles de tolerancia a través de diversos mecanismos, las vías de señalización también pueden diferir, por lo que no se cuenta con un modelo único. Algunas de las moléculas comunes que participan en la respuesta de tolerancia se determinaron. Para identificar genes candidatos a utilizar en el desarrollo de cultivares tolerantes al aluminio, se consultó artículos científicos publicados entre 1987 y 2013. Mediante el reporte de uso de técnicas convencionales de hibridación, mutación, selección asistida por marcadores moleculares y transferencia de genes, se obtuvieron datos de materiales tolerantes evidenciándose mecanismos moleculares para tolerancia al aluminio. Se encontraron reportados genes total y parcialmente caracterizados, con uso potencial en ingeniería genética y en selección asistida por marcadores, para la obtención de genotipos tolerantes al aluminio.
Agricultural production in the Colombian Orinoco is affected by the high aluminum content found in 4.5 million hectares. Genotypes of different species have acquired different levels of tolerance and signaling pathways through various mechanisms, making a single model impossible. Some of the molecules commonly involved in the tolerance response have already been identified. To identify candidate genes to produce aluminum-tolerant cultivars, we consulted scientific articles published between 1987 and 2013. We obtained data of aluminum-tolerant materials and molecular mechanisms for tolerance through reports of techniques using hybridization, mutation, molecular marker-assisted selection and gene transfer. We found several reports on wholly or partially characterized genes with potential use in genetic engineering and in marker assisted selection to obtain aluminum tolerant genotypes.
Um problema para a produção agrícola no Orinoco da Colômbia, são os 4,5 milhões de hectares com alto teor de alumínio. Genótipos de diferentes espécies apresentam tolerância através de diferentes mecanismos, as vias de sinalização também podem ser diferentes, de modo que não existe um modelo único. Algumas das moléculas comuns envolvidas na resposta da tolerância foram determinadas. A fim de identificar genes candidatos para uso no desenvolvimento de cultivos tolerantes ao alumínio foram consultados artigos científicos publicados entre 1987 e 2013. Por meio de técnicas convencionais que utilizam a hibridização, mutação, seleção assistida por marcadores moleculares e de transferência de genes, obtiveram-se dados de materiais tolerantes e se evidenciou mecanismos moleculares para tolerância ao alumínio. Encontraram-se reportados vários genes total e parcialmente caracterizados, com uso potencial na engenharia genética e na seleção assistida por marcadores, para a obtenção de genótipos tolerantes ao alumínio.
RESUMO
Impaired root development caused by aluminum (Al) toxicity is a major cause of grain yield reduction in crops cultivated on acid soils, which are widespread worldwide. In sorghum, the major Al-tolerance locus, AltSB , is due to the function of SbMATE, which is an Al-activated root citrate transporter. Here we performed a molecular and physiological characterization of various AltSB donors and near-isogenic lines harboring various AltSB alleles. We observed a partial transfer of Al tolerance from the parents to the near-isogenic lines that was consistent across donor alleles, emphasizing the occurrence of strong genetic background effects related to AltSB . This reduction in tolerance was variable, with a 20% reduction being observed when highly Al-tolerant lines were the AltSB donors, and a reduction as great as 70% when other AltSB alleles were introgressed. This reduction in Al tolerance was closely correlated with a reduction in SbMATE expression in near-isogenic lines, suggesting incomplete transfer of loci acting in trans on SbMATE. Nevertheless, AltSB alleles from the highly Al-tolerant sources SC283 and SC566 were found to retain high SbMATE expression, presumably via elements present within or near the AltSB locus, resulting in significant transfer of the Al-tolerance phenotype to the derived near-isogenic lines. Allelic effects could not be explained by coding region polymorphisms, although occasional mutations may affect Al tolerance. Finally, we report on the extensive occurrence of alternative splicing for SbMATE, which may be an important component regulating SbMATE expression in sorghum by means of the nonsense-mediated RNA decay pathway.
Assuntos
Alumínio/toxicidade , Regulação da Expressão Gênica de Plantas/efeitos dos fármacos , Proteínas de Plantas/metabolismo , Sorghum/efeitos dos fármacos , Sorghum/metabolismo , Processamento Alternativo , Sequência de Bases , Variação Genética , Genoma de Planta , Modelos Moleculares , Dados de Sequência Molecular , Proteínas de Plantas/genética , Conformação Proteica , Sorghum/genéticaRESUMO
Medicine plants play an important role in public health, especially in developing countries, and among them Cunila galioides Benth. is a naturally occurring medicinal and aromatic plant native from south Brazil. Flavonoids, present in significant amounts in some species of Cunila sp., possess several properties and its content may be affected by aluminum stress. In this context, the aim of this work was to assess the influence of different levels of aluminum in nutrient solution on the growth and content of essential oils and flavonoids in three chemotypes of Cunila galioides. The plant parameters evaluated were aerial dry matter, dry root matter, root length, root score, calcium and aluminum concentrations in the vegetable tissue, flavonoids content as well as essential oil yield and its major components concentration. Results showed that the addition of 30.0 mg.L-¹ of Al³+ to the nutrient solution leads to a decrease of all agronomic parameters evaluated with losses in the cultivation yield for the greatest concentration studied. The André da Rocha population was found to be the most tolerant to aluminum while Bom Jardim da Serra was the most sensitive. The aluminum concentration did not influence the extraction yield as well as the content of major components in the oils. The concentration of flavonoids increased significantly for the populations tolerant to aluminum concentration in the nutrient solution, thus indicating a relationship between tolerance and flavonoids production, which may be an interesting characteristic from the medicinal point of view.
Plantas medicinais desempenham um papel importante na saúde pública, especialmente em países em desenvolvimento, e entre elas Cunila galioides Benth., que é uma planta nativa de ocorrência no sul do Brasil. Os flavonoides, presentes em quantidades significativas em algumas espécies de Cunila sp., possuem diversas propriedades, sendo que seu teor pode ser afetado por estresse de alumínio. Neste contexto, o objetivo deste trabalho foi avaliar a influência de diferentes níveis de alumínio na solução nutritiva, em relação ao crescimento e teor de óleos essenciais e de flavonoides, em três quimiotipos de Cunila galioides Benth. Os parâmetros avaliados foram matéria seca da parte aérea, matéria seca da raiz, comprimento da raiz, contagem de raízes, concentrações de cálcio e alumínio no tecido vegetal, teor de flavonoides, assim como o rendimento do óleo essencial e concentração dos seus componentes principais. Os resultados mostraram que a adição de 30,0 mg.L-¹ de Al³+ à solução nutriente conduz à diminuição de todos os parâmetros agronômicos avaliados com perdas de rendimento em cultivos com a maior concentração avaliada. A população de André da Rocha foi a mais tolerante ao alumínio, enquanto que Bom Jardim da Serra foi a mais sensível. A concentração de alumínio não influenciou no rendimento de extração, assim como no teor dos componentes principais nos óleos. A concentração de flavonoides aumentou significativamente nas populações tolerantes ao alumínio, indicando uma relação entre a tolerância ao Al³+ e a produção de flavonoides, o que pode ser interessante do ponto de vista medicinal.
Assuntos
Plantas Medicinais , Flavonoides/biossíntese , Óleos Voláteis , Lamiaceae/química , Alumínio/farmacologiaRESUMO
Medicine plants play an important role in public health, especially in developing countries, and among them Cunila galioides Benth. is a naturally occurring medicinal and aromatic plant native from south Brazil. Flavonoids, present in significant amounts in some species of Cunila sp., possess several properties and its content may be affected by aluminum stress. In this context, the aim of this work was to assess the influence of different levels of aluminum in nutrient solution on the growth and content of essential oils and flavonoids in three chemotypes of Cunila galioides. The plant parameters evaluated were aerial dry matter, dry root matter, root length, root score, calcium and aluminum concentrations in the vegetable tissue, flavonoids content as well as essential oil yield and its major components concentration. Results showed that the addition of 30.0 mg.L--1 of Al3+ to the nutrient solution leads to a decrease of all agronomic parameters evaluated with losses in the cultivation yield for the greatest concentration studied. The André da Rocha population was found to be the most tolerant to aluminum while Bom Jardim da Serra was the most sensitive. The aluminum concentration did not influence the extraction yield as well as the content of major components in the oils. The concentration of flavonoids increased significantly for the populations tolerant to aluminum concentration in the nutrient solution, thus indicating a relationship between tolerance and flavonoids production, which may be an interesting characteristic from the medicinal point of view.
Plantas medicinais desempenham um papel importante na saúde pública, especialmente em países em desenvolvimento, e entre elas Cunila galioides Benth., que é uma planta nativa de ocorrência no sul do Brasil. Os flavonoides, presentes em quantidades significativas em algumas espécies de Cunila sp., possuem diversas propriedades, sendo que seu teor pode ser afetado por estresse de alumínio. Neste contexto, o objetivo deste trabalho foi avaliar a influência de diferentes níveis de alumínio na solução nutritiva, em relação ao crescimento e teor de óleos essenciais e de flavonoides, em três quimiotipos de Cunila galioides Benth. Os parâmetros avaliados foram matéria seca da parte aérea, matéria seca da raiz, comprimento da raiz, contagem de raízes, concentrações de cálcio e alumínio no tecido vegetal, teor de flavonoides, assim como o rendimento do óleo essencial e concentração dos seus componentes principais. Os resultados mostraram que a adição de 30,0 mg.L-1 de Al3+ à solução nutriente conduz à diminuição de todos os parâmetros agronômicos avaliados com perdas de rendimento em cultivos com a maior concentração avaliada. A população de André da Rocha foi a mais tolerante ao alumínio, enquanto que Bom Jardim da Serra foi a mais sensível. A concentração de alumínio não influenciou no rendimento de extração, assim como no teor dos componentes principais nos óleos. A concentração de flavonoides aumentou significativamente nas populações tolerantes ao alumínio, indicando uma relação entre a tolerância ao Al3+ e a produção de flavonoides, o que pode ser interessante do ponto de vista medicinal.
RESUMO
Aluminum (Al) toxicity is the main factor limiting crop productivity in acidic soils around the world. In cereals, this problem reduces crop yields by 30-40 percent. The use of DNA-based markers linked to phenotypic traits is an interesting alternative approach. Strategies such as molecular marker-assisted selection (MAS) in conjunction with bioinformatics-based tools such as graphical genotypes (GGT) have been important for confirming introgression of genes or genomic regions in cereals but also to reduce the time and cost of identifying them through genetic selection. These biotechnologies also make it possible to identify target genes or quantitative trait loci (QTL) that can be potentially used in similar crops to increase their productivity. This review presents the main advances in the genetic improvement of cereals for Al-tolerance.
Assuntos
Alumínio/toxicidade , Mapeamento Cromossômico , Grão Comestível/genética , Transportadores de Ânions Orgânicos , Resistência a Medicamentos/genética , Produção Agrícola , Grão Comestível , Cruzamento/métodos , Marcadores Genéticos , Ácidos Orgânicos , Locos de Características Quantitativas , Seleção GenéticaRESUMO
In Brazil, acid soils represent a large portion of the agricultural area and the development of cultivars with tolerance to acidity has shown to be feasible. The identification of germoplasm potentially tolerant to acidity was the purpose of this work. Two distinct groups of maize (Zea mays L.) populations were crossed according to the partial diallel mating scheme: Group [1] with six adapted populations and Group [2] with seven exotic varieties. All varieties and crosses were evaluated under the condition of acid and low fertility soil. Traits analyzed were: EY- ear yield, PH- plant height and EH- ear height. Outstanding varieties per se, with yield higher than 3 t ha-1, were SUWAN-2, TAITINGA, TAIÚBA and IUBATÃ in Group [1]; and CUPURICO DMR, TUXPEÑO AMARILLO, and AMARILLO DENTADO in Group [2]. Heterosis and its components showed no significant variation for PH and EH; and only average heterosis was significant for EY. The general combining ability effects (g i and g j) showed significant variation (P 0.01) for all traits and were due mainly to the variation of variety effects. The higher estimates g i for EY were exhibited by WP 12 (0.178 t ha-1), TAIÚBA (0.176) and TAITINGA (0.161) in Group [1]; and for g j by CUPURICO DMR (0.300), TAITINGA TUXPEÑO AMARILLO (0.280) in Group [2], respectively. The variety cross TAITINGA x TUXPEÑO AMARILLO with mean yield of 4.34 t ha-1 and 25% of mid-parent heterosis may be indicated as an heterotic pattern. PH and EH were not considered as limiting to preclude the use of the outstanding populations. The varieties SUWAN 2, CUPURICO DMR, and TAITINGA were suggested to be used in maize breeding programs for acid soils.
No Brasil os solos ácidos representam uma grande parte da área agricultável e o desenvolvimento de cultivares com tolerância à acidez tem-se mostrado exequível. A identificação de germoplasma potencialmente tolerante à acidez foi o propósito deste trabalho. Dois grupos distintos de populações de milho (Zea mays L.) foram cruzados segundo o esquema de dialelo parcial: Grupo [1] com seis populações adaptadas e Grupo [2] com sete exóticas. As populações e seus híbridos foram avaliados sob condição de solo ácido e de baixa fertilidade. Os caracteres analisados foram: EY- peso de espigas, PH- altura da planta e EH- altura da espiga. As populações que sobressaíram, com produtividade per se acima de 3 t ha-1 foram SUWAN-2, TAITINGA, TAIÚBA e IUBATÃ no Grupo [1]; e CUPURICO DMR, TUXPEÑO AMARILLO, e AMARILLO DENTADO no Grupo [2]. A heterose e seus componentes mostraram variação não significativa para PH e EH; e somente a heterose média foi significativa para EY. Os efeitos de capacidade geral de combinação (g i e g j) mostraram variação significativa (P 0,01) para todos os caracteres e foram devidos principalmente à variação dos efeitos de variedades. As maiores estimativas de g i no Grupo [1] para EY (t ha-1) foram exibidas por WP 12 (0,178 t ha-1), TAIÚBA (0,176) and TAITINGA (0.161) no Grupo [1]; e para g j por CUPURICO DMR (0.300), TAITINGA TUXPEÑO AMARILLO (0.280) no Grupo [2], respectivamente. O híbrido interpopulacional TAITINGA x TUXPEÑO AMARILLO com produção média de 4,34 t ha-1 e 25% de heterose pode ser indicado como um padrão heterótico. A altura da planta e da espiga não foram consideradas limitantes para impedir o uso das populações mais promissoras. As variedades SUWAN 2, CUPURICO DMR e TAITINGA foram as mais promissoras para programas de mehoramento visando adaptação às condições de solo ácidos.
RESUMO
Acid soils and associated mineral toxicity are a great restriction for crop production, and among the metals, aluminum is considered one of the most important problems in soils with pH £ 5.0. Breeding for aluminum tolerance is considered a useful way to improve important economic plants' performance in acid soils with high concentration of this metal, as those which occur in South Brazil. However, for this, it is necessary to identify aluminum-tolerant genotypes, to understand aluminum phytotoxicity and tolerance mechanisms, and to know genetic control of this tolerance. This paper presents a review of these subjects, with emphasis on aluminum forms present in soils, their effect over the plants, the stage of knowledge on aluminum tolerance mechanisms and their genetic control in different cereal groups.
Solos ácidos associados a metais tóxicos são uma das maiores restrições à produção agrícola e, entre os metais, o alumínio destaca-se como um dos maiores problemas em solos com pH £ 5,0. O melhoramento para tolerância ao alumínio é considerado um excelente caminho para aumentar o desempenho de plantas economicamente importantes cultivadas em solos ácidos ricos nesse metal, como os que ocorrem no sul do Brasil. Para isso, é necessário identificar, entre os diferentes cultivares, aqueles que apresentam mecanismos de tolerância, compreender como esses cultivares agem em sua presença, bem como conhecer o controle genético da tolerância ao alumínio. Este trabalho apresenta uma revisão sobre o assunto, abordando as formas de alumínio presentes no solo, seu efeito sobre as plantas, os mecanismos de tolerância e o controle genético desta tolerância em diferentes grupos de cereais.
RESUMO
In Brazil, acid soils represent a large portion of the agricultural area and the development of cultivars with tolerance to acidity has shown to be feasible. The identification of germoplasm potentially tolerant to acidity was the purpose of this work. Two distinct groups of maize (Zea mays L.) populations were crossed according to the partial diallel mating scheme: Group [1] with six adapted populations and Group [2] with seven exotic varieties. All varieties and crosses were evaluated under the condition of acid and low fertility soil. Traits analyzed were: EY- ear yield, PH- plant height and EH- ear height. Outstanding varieties per se, with yield higher than 3 t ha-1, were SUWAN-2, TAITINGA, TAIÚBA and IUBATÃ in Group [1]; and CUPURICO DMR, TUXPEÑO AMARILLO, and AMARILLO DENTADO in Group [2]. Heterosis and its components showed no significant variation for PH and EH; and only average heterosis was significant for EY. The general combining ability effects (g i and g j) showed significant variation (P 0.01) for all traits and were due mainly to the variation of variety effects. The higher estimates g i for EY were exhibited by WP 12 (0.178 t ha-1), TAIÚBA (0.176) and TAITINGA (0.161) in Group [1]; and for g j by CUPURICO DMR (0.300), TAITINGA TUXPEÑO AMARILLO (0.280) in Group [2], respectively. The variety cross TAITINGA x TUXPEÑO AMARILLO with mean yield of 4.34 t ha-1 and 25% of mid-parent heterosis may be indicated as an heterotic pattern. PH and EH were not considered as limiting to preclude the use of the outstanding populations. The varieties SUWAN 2, CUPURICO DMR, and TAITINGA were suggested to be used in maize breeding programs for acid soils.
No Brasil os solos ácidos representam uma grande parte da área agricultável e o desenvolvimento de cultivares com tolerância à acidez tem-se mostrado exequível. A identificação de germoplasma potencialmente tolerante à acidez foi o propósito deste trabalho. Dois grupos distintos de populações de milho (Zea mays L.) foram cruzados segundo o esquema de dialelo parcial: Grupo [1] com seis populações adaptadas e Grupo [2] com sete exóticas. As populações e seus híbridos foram avaliados sob condição de solo ácido e de baixa fertilidade. Os caracteres analisados foram: EY- peso de espigas, PH- altura da planta e EH- altura da espiga. As populações que sobressaíram, com produtividade per se acima de 3 t ha-1 foram SUWAN-2, TAITINGA, TAIÚBA e IUBATÃ no Grupo [1]; e CUPURICO DMR, TUXPEÑO AMARILLO, e AMARILLO DENTADO no Grupo [2]. A heterose e seus componentes mostraram variação não significativa para PH e EH; e somente a heterose média foi significativa para EY. Os efeitos de capacidade geral de combinação (g i e g j) mostraram variação significativa (P 0,01) para todos os caracteres e foram devidos principalmente à variação dos efeitos de variedades. As maiores estimativas de g i no Grupo [1] para EY (t ha-1) foram exibidas por WP 12 (0,178 t ha-1), TAIÚBA (0,176) and TAITINGA (0.161) no Grupo [1]; e para g j por CUPURICO DMR (0.300), TAITINGA TUXPEÑO AMARILLO (0.280) no Grupo [2], respectivamente. O híbrido interpopulacional TAITINGA x TUXPEÑO AMARILLO com produção média de 4,34 t ha-1 e 25% de heterose pode ser indicado como um padrão heterótico. A altura da planta e da espiga não foram consideradas limitantes para impedir o uso das populações mais promissoras. As variedades SUWAN 2, CUPURICO DMR e TAITINGA foram as mais promissoras para programas de mehoramento visando adaptação às condições de solo ácidos.
RESUMO
Acid soils and associated mineral toxicity are a great restriction for crop production, and among the metals, aluminum is considered one of the most important problems in soils with pH £ 5.0. Breeding for aluminum tolerance is considered a useful way to improve important economic plants' performance in acid soils with high concentration of this metal, as those which occur in South Brazil. However, for this, it is necessary to identify aluminum-tolerant genotypes, to understand aluminum phytotoxicity and tolerance mechanisms, and to know genetic control of this tolerance. This paper presents a review of these subjects, with emphasis on aluminum forms present in soils, their effect over the plants, the stage of knowledge on aluminum tolerance mechanisms and their genetic control in different cereal groups.
Solos ácidos associados a metais tóxicos são uma das maiores restrições à produção agrícola e, entre os metais, o alumínio destaca-se como um dos maiores problemas em solos com pH £ 5,0. O melhoramento para tolerância ao alumínio é considerado um excelente caminho para aumentar o desempenho de plantas economicamente importantes cultivadas em solos ácidos ricos nesse metal, como os que ocorrem no sul do Brasil. Para isso, é necessário identificar, entre os diferentes cultivares, aqueles que apresentam mecanismos de tolerância, compreender como esses cultivares agem em sua presença, bem como conhecer o controle genético da tolerância ao alumínio. Este trabalho apresenta uma revisão sobre o assunto, abordando as formas de alumínio presentes no solo, seu efeito sobre as plantas, os mecanismos de tolerância e o controle genético desta tolerância em diferentes grupos de cereais.
RESUMO
Diallel crosses were made involving the cultivars: 'BH-1146', aluminum tolerant; 'IAC-161', moderately aluminum tolerant, and 'Anahuac', aluminum sensitive. Parents, F1s, F2s and reciprocal backcrosses were tested for their seedling reaction to 6 mg/L of Al3+ in nutrient solution, in laboratory conditions, and evaluated for grain yield, plant height, head length and rachis internode length at maturity, in pots, under screen house conditions in Campinas, SP, Brazil. Narrow sense heritability estimates were moderate to high for plant height (0.638 - 0.860), moderate for aluminum tolerance (0.409 - 0.593) and head length (0.390 - 0.522) except for the hybrid 'Anahuac' x 'IAC-161', which presented a high value (0.851), and low to moderate for grain yield (0.054 - 0.463) and rachis internode length (0.090 - 0.595). Phenotypic correlations between tolerance to Al3+ and plant height, rachis internode length and grain yield were only significant for one of the hybrids. Phenotypic correlations between plant height with grain yield and head length, and between head length with rachis internode length were positive and significant for all crosses, showing association between these characters. Results suggest that it would be possible to select desirable plants from eventual recombinations if large segregating populations are used.
Visando estimar a herdabilidade em sentido restrito para a tolerância ao Al3+, altura das plantas, comprimento da espiga, comprimento do internódio da raque e produção de grãos, bem como as correlações entre essas características, foram efetuados cruzamentos, em forma dialélica entre os cultivares 'BH-1146', tolerante ao Al3+ e de porte alto; 'IAC-161' e 'Anahuac' ambos de porte semi-anão, com média tolerância e sensibilidade à toxicidade de Al3+, respectivamente. Plântulas representando os pais, as gerações F1 e F2 e os retrocruzamentos para ambos os pais, foram testadas para a reação a 6 mg/L de Al3+ em solução nutritiva. As plantas, devidamente identificadas, foram transplantadas para vasos localizados no telado. Os valores da herdabilidade em sentido restrito para a tolerância ao Al3+ foram médios (0,409 a 0,593) e para altura de planta foram médios a altos (0,638 a 0,860) para os três híbridos avaliados. Em relação ao comprimento da espiga, os valores foram médios para os híbridos 'BH-1146' x 'IAC-161' (0,390) e 'BH-1146' x 'Anahuac' (0,522) e alto para Anahuac x IAC-161 (0,851). Valores baixos a médios da herdabilidade em sentido restrito foram estimados para os caracteres comprimento do internódio da raque (0,090 a 0,595) e produção de grãos (0,054 a 0,463). As correlações fenotípicas entre a tolerância ao Al3+ somente foram associadas significativamente com altura de planta, comprimento do internódio da raque e produção de grãos em um dos híbridos estudados e as correlações fenotípicas entre altura da planta com produção de grãos e comprimento da espiga, e entre comprimento da espiga e comprimento do internódio da raque foram positivas e significativas em todos os cruzamentos, mostrando haver associação entre esses caracteres, portanto, o estudo de grandes populações F2 seria de interesse para proporcionar maior freqüência de recombinantes desejáveis.