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1.
Mol Ecol ; 28(15): 3544-3560, 2019 08.
Artigo em Inglês | MEDLINE | ID: mdl-31287919

RESUMO

Understanding the genomic basis of adaptation in maize is important for gene discovery and the improvement of breeding germplasm, but much remains a mystery in spite of significant population genetics and archaeological research. Identifying the signals underpinning adaptation are challenging as adaptation often coincided with genetic drift, and the base genomic diversity of the species in massive. In this study, tGBS technology was used to genotype 1,143 diverse maize accessions including landraces collected from 20 countries and elite breeding lines of tropical lowland, highland, subtropical/midaltitude and temperate ecological zones. Based on 355,442 high-quality single nucleotide polymorphisms, 13 genomic regions were detected as being under selection using the bottom-up searching strategy, EigenGWAS. Of the 13 selection regions, 10 were first reported, two were associated with environmental parameters via EnvGWAS, and 146 genes were enriched. Combining large-scale genomic and ecological data in this diverse maize panel, our study supports a polygenic adaptation model of maize and offers a framework to enhance our understanding of both the mechanistic basis and the evolutionary consequences of maize domestication and adaptation. The regions identified here are promising candidates for further, targeted exploration to identify beneficial alleles and haplotypes for deployment in maize breeding.

2.
PLoS One ; 11(6): e0157236, 2016.
Artigo em Inglês | MEDLINE | ID: mdl-27280295

RESUMO

Quality control (QC) of germplasm identity and purity is a critical component of breeding and conservation activities. SNP genotyping technologies and increased availability of markers provide the opportunity to employ genotyping as a low-cost and robust component of this QC. In the public sector available low-cost SNP QC genotyping methods have been developed from a very limited panel of markers of 1,000 to 1,500 markers without broad selection of the most informative SNPs. Selection of optimal SNPs and definition of appropriate germplasm sampling in addition to platform section impact on logistical and resource-use considerations for breeding and conservation applications when mainstreaming QC. In order to address these issues, we evaluated the selection and use of SNPs for QC applications from large DArTSeq data sets generated from CIMMYT maize inbred lines (CMLs). Two QC genotyping strategies were developed, the first is a "rapid QC", employing a small number of SNPs to identify potential mislabeling of seed packages or plots, the second is a "broad QC", employing a larger number of SNP, used to identify each germplasm entry and to measure heterogeneity. The optimal marker selection strategies combined the selection of markers with high minor allele frequency, sampling of clustered SNP in proportion to marker cluster distance and selecting markers that maintain a uniform genomic distribution. The rapid and broad QC SNP panels selected using this approach were further validated using blind test assessments of related re-generation samples. The influence of sampling within each line was evaluated. Sampling 192 individuals would result in close to 100% possibility of detecting a 5% contamination in the entry, and approximately a 98% probability to detect a 2% contamination of the line. These results provide a framework for the establishment of QC genotyping. A comparison of financial and time costs for use of these approaches across different platforms is discussed providing a framework for institutions involved in maize conservation and breeding to assess the resource use effectiveness of QC genotyping. Application of these research findings, in combination with existing QC approaches, will ensure the regeneration, distribution and use in breeding of true to type inbred germplasm. These findings also provide an effective approach to optimize SNP selection for QC genotyping in other species.


Assuntos
Técnicas de Genotipagem/métodos , Polimorfismo de Nucleotídeo Único , Zea mays/genética , Marcadores Genéticos , Controle de Qualidade
3.
Theor Appl Genet ; 129(4): 753-765, 2016 Apr.
Artigo em Inglês | MEDLINE | ID: mdl-26849239

RESUMO

KEY MESSAGE: Molecular characterization information on genetic diversity, population structure and genetic relationships provided by this research will help maize breeders to better understand how to utilize the current CML collection. CIMMYT maize inbred lines (CMLs) have been widely used all over the world and have contributed greatly to both tropical and temperate maize improvement. Genetic diversity and population structure of the current CML collection and of six temperate inbred lines were assessed and relationships among all lines were determined with genotyping-by-sequencing SNPs. Results indicated that: (1) wider genetic distance and low kinship coefficients among most pairs of lines reflected the uniqueness of most lines in the current CML collection; (2) the population structure and genetic divergence between the Temperate subgroup and Tropical subgroups were clear; three major environmental adaptation groups (Lowland Tropical, Subtropical/Mid-altitude and Highland Tropical subgroups) were clearly present in the current CML collection; (3) the genetic diversity of the three Tropical subgroups was similar and greater than that of the Temperate subgroup; the average genetic distance between the Temperate and Tropical subgroups was greater than among Tropical subgroups; and (4) heterotic patterns in each environmental adaptation group estimated using GBS SNPs were only partially consistent with patterns estimated based on combining ability tests and pedigree information. Combining current heterotic information based on combining ability tests and the genetic relationships inferred from molecular marker analyses may be the best strategy to define heterotic groups for future tropical maize improvement. Information resulting from this research will help breeders to better understand how to utilize all the CMLs to select parental lines, replace testers, assign heterotic groups and create a core set of breeding germplasm.


Assuntos
Genótipo , Vigor Híbrido , Polimorfismo de Nucleotídeo Único , Zea mays/genética , DNA de Plantas/genética , Frequência do Gene , Endogamia , Melhoramento Vegetal , Análise de Sequência de DNA
4.
G3 (Bethesda) ; 5(5): 891-909, 2015 Mar 12.
Artigo em Inglês | MEDLINE | ID: mdl-25770100

RESUMO

Genotyping by sequencing allows for large-scale genetic analyses in plant species with no reference genome, but sets the challenge of sound inference in presence of uncertain genotypes. We report an imputation-based genome-wide association study (GWAS) in reed canarygrass (Phalaris arundinacea L., Phalaris caesia Nees), a cool-season grass species with potential as a biofuel crop. Our study involved two linkage populations and an association panel of 590 reed canarygrass genotypes. Plants were assayed for up to 5228 single nucleotide polymorphism markers and 35 traits. The genotypic markers were derived from low-depth sequencing with 78% missing data on average. To soundly infer marker-trait associations, multiple imputation (MI) was used: several imputes of the marker data were generated to reflect imputation uncertainty and association tests were performed on marker effects across imputes. A total of nine significant markers were identified, three of which showed significant homology with the Brachypodium dystachion genome. Because no physical map of the reed canarygrass genome was available, imputation was conducted using classification trees. In general, MI showed good consistency with the complete-case analysis and adequate control over imputation uncertainty. A gain in significance of marker effects was achieved through MI, but only for rare cases when missing data were <45%. In addition to providing insight into the genetic basis of important traits in reed canarygrass, this study presents one of the first applications of MI to genome-wide analyses and provides useful guidelines for conducting GWAS based on genotyping-by-sequencing data.


Assuntos
Genoma de Planta , Estudo de Associação Genômica Ampla , Sequenciamento de Nucleotídeos em Larga Escala , Característica Quantitativa Herdável , Algoritmos , Biocombustíveis , Modelos Teóricos
5.
PLoS One ; 9(11): e112227, 2014.
Artigo em Inglês | MEDLINE | ID: mdl-25390940

RESUMO

Switchgrass (Panicum virgatum L.) is a perennial grass undergoing development as a biofuel feedstock. One of the most important factors hindering breeding efforts in this species is the need for accurate measurement of biomass yield on a per-hectare basis. Genomic selection on simple-to-measure traits that approximate biomass yield has the potential to significantly speed up the breeding cycle. Recent advances in switchgrass genomic and phenotypic resources are now making it possible to evaluate the potential of genomic selection of such traits. We leveraged these resources to study the ability of three widely-used genomic selection models to predict phenotypic values of morphological and biomass quality traits in an association panel consisting of predominantly northern adapted upland germplasm. High prediction accuracies were obtained for most of the traits, with standability having the highest ten-fold cross validation prediction accuracy (0.52). Moreover, the morphological traits generally had higher prediction accuracies than the biomass quality traits. Nevertheless, our results suggest that the quality of current genomic and phenotypic resources available for switchgrass is sufficiently high for genomic selection to significantly impact breeding efforts for biomass yield.


Assuntos
Genoma de Planta , Panicum/crescimento & desenvolvimento , Panicum/genética , Biocombustíveis , Biomassa , Ecótipo , Fenótipo , Ploidias , Polimorfismo de Nucleotídeo Único , Análise de Componente Principal , Seleção Genética/genética , Espectroscopia de Luz Próxima ao Infravermelho
6.
PLoS Genet ; 9(1): e1003215, 2013.
Artigo em Inglês | MEDLINE | ID: mdl-23349638

RESUMO

Switchgrass (Panicum virgatum L.) is a perennial grass that has been designated as an herbaceous model biofuel crop for the United States of America. To facilitate accelerated breeding programs of switchgrass, we developed both an association panel and linkage populations for genome-wide association study (GWAS) and genomic selection (GS). All of the 840 individuals were then genotyped using genotyping by sequencing (GBS), generating 350 GB of sequence in total. As a highly heterozygous polyploid (tetraploid and octoploid) species lacking a reference genome, switchgrass is highly intractable with earlier methodologies of single nucleotide polymorphism (SNP) discovery. To access the genetic diversity of species like switchgrass, we developed a SNP discovery pipeline based on a network approach called the Universal Network-Enabled Analysis Kit (UNEAK). Complexities that hinder single nucleotide polymorphism discovery, such as repeats, paralogs, and sequencing errors, are easily resolved with UNEAK. Here, 1.2 million putative SNPs were discovered in a diverse collection of primarily upland, northern-adapted switchgrass populations. Further analysis of this data set revealed the fundamentally diploid nature of tetraploid switchgrass. Taking advantage of the high conservation of genome structure between switchgrass and foxtail millet (Setaria italica (L.) P. Beauv.), two parent-specific, synteny-based, ultra high-density linkage maps containing a total of 88,217 SNPs were constructed. Also, our results showed clear patterns of isolation-by-distance and isolation-by-ploidy in natural populations of switchgrass. Phylogenetic analysis supported a general south-to-north migration path of switchgrass. In addition, this analysis suggested that upland tetraploid arose from upland octoploid. All together, this study provides unparalleled insights into the diversity, genomic complexity, population structure, phylogeny, phylogeography, ploidy, and evolutionary dynamics of switchgrass.


Assuntos
Variação Genética , Estudo de Associação Genômica Ampla , Panicum/genética , Poliploidia , Biocombustíveis , Evolução Biológica , Mapeamento Cromossômico , Genoma de Planta , Filogenia , Filogeografia , Polimorfismo de Nucleotídeo Único , Seleção Genética , Análise de Sequência de DNA , Sintenia
7.
PLoS One ; 7(9): e44112, 2012.
Artigo em Inglês | MEDLINE | ID: mdl-23049744

RESUMO

Although yield trials for switchgrass (Panicum virgatum L.), a potentially high value biofuel feedstock crop, are currently underway throughout North America, the genetic tools for crop improvement in this species are still in the early stages of development. Identification of high-density molecular markers, such as single nucleotide polymorphisms (SNPs), that are amenable to high-throughput genotyping approaches, is the first step in a quantitative genetics study of this model biofuel crop species. We generated and sequenced expressed sequence tag (EST) libraries from thirteen diverse switchgrass cultivars representing both upland and lowland ecotypes, as well as tetraploid and octoploid genomes. We followed this with reduced genomic library preparation and massively parallel sequencing of the same samples using the Illumina Genome Analyzer technology platform. EST libraries were used to generate unigene clusters and establish a gene-space reference sequence, thus providing a framework for assembly of the short sequence reads. SNPs were identified utilizing these scaffolds. We used a custom software program for alignment and SNP detection and identified over 149,000 SNPs across the 13 short-read sequencing libraries (SRSLs). Approximately 25,000 additional SNPs were identified from the entire EST collection available for the species. This sequencing effort generated data that are suitable for marker development and for estimation of population genetic parameters, such as nucleotide diversity and linkage disequilibrium. Based on these data, we assessed the feasibility of genome wide association mapping and genomic selection applications in switchgrass. Overall, the SNP markers discovered in this study will help facilitate quantitative genetics experiments and greatly enhance breeding efforts that target improvement of key biofuel traits and development of new switchgrass cultivars.


Assuntos
Etiquetas de Sequências Expressas , Genoma de Planta , Biblioteca Genômica , Panicum/genética , Polimorfismo de Nucleotídeo Único , Biocombustíveis , Mapeamento Cromossômico , Ecótipo , Marcadores Genéticos , Estudo de Associação Genômica Ampla , Genótipo , Sequenciamento de Nucleotídeos em Larga Escala , América do Norte , Panicum/classificação , Filogeografia , Ploidias
8.
Plant Cell ; 21(8): 2194-202, 2009 Aug.
Artigo em Inglês | MEDLINE | ID: mdl-19654263

RESUMO

The goal of many plant scientists' research is to explain natural phenotypic variation in terms of simple changes in DNA sequence. Traditionally, linkage mapping has been the most commonly employed method to reach this goal: experimental crosses are made to generate a family with known relatedness, and attempts are made to identify cosegregation of genetic markers and phenotypes within this family. In vertebrate systems, association mapping (also known as linkage disequilibrium mapping) is increasingly being adopted as the mapping method of choice. Association mapping involves searching for genotype-phenotype correlations in unrelated individuals and often is more rapid and cost-effective than traditional linkage mapping. We emphasize here that linkage and association mapping are complementary approaches and are more similar than is often assumed. Unlike in vertebrates, where controlled crosses can be expensive or impossible (e.g., in humans), the plant scientific community can exploit the advantages of both controlled crosses and association mapping to increase statistical power and mapping resolution. While the time and money required for the collection of genotype data were critical considerations in the past, the increasing availability of inexpensive DNA sequencing and genotyping methods should prompt researchers to shift their attention to experimental design. This review provides thoughts on finding the optimal experimental mix of association mapping using unrelated individuals and controlled crosses to identify the genes underlying phenotypic variation.


Assuntos
Mapeamento Cromossômico/métodos , Projetos de Pesquisa , Animais , Genótipo , Humanos , Fenótipo
9.
Genet Res (Camb) ; 90(1): 111-8, 2008 Feb.
Artigo em Inglês | MEDLINE | ID: mdl-18289405

RESUMO

One of the long-standing mysteries in genomic evolution is the observation that much of the genome is composed of repetitive DNA, resulting in inter- and intraspecific variation in nuclear DNA content. Our discovery of a negative correlation between nuclear DNA content and flower size in Silene latifolia has been supported by our subsequent investigation of changes in DNA content as a correlated response to selection on flower size. Moreover, we have observed a similar trend across a range of related dioecious species in Silene sect. Elisanthe. Given the presence of sex chromosomes in dioecious Silene species, and the tendency of sex chromosomes to accumulate repetitive DNA, it seems plausible that dioecious species undergo genomic evolution in ways that differ from what one might expect in hermaphroditic species. Specifically, we query whether the observed relationship between nuclear DNA content and flower size observed in dioecious Silene is a peculiarity of sex chromosome evolution. In the present study we investigated nuclear DNA content and flower size variation in hermaphroditic species of Silene sect. Siphonomorpha, as close relatives of the dioecious species studied previously. Although the nuclear DNA contents of these species were lower than those for species in sect. Elisanthe, there was still significant intra- as well as interspecific variation in nuclear DNA content. Flower size variation was found among species of sect. Siphonomorpha for petal claw and petal limb lengths, but not for calyx diameter. This last trait varies extensively in sect. Elisanthe, in part due to sex-specific selection. A negative correlation with nuclear DNA content was found across populations for petal limb length, but not for other floral dimensions. We conclude that impacts of nuclear DNA content on phenotypic evolution do manifest themselves in hermaphroditic species, so that the effects observed in sect. Elisanthe, and particularly in S. latifolia, while perhaps amplified by the genomic impacts of sex chromosomes, are not limited to dioecious taxa.


Assuntos
Evolução Biológica , DNA Intergênico/genética , Fenótipo , Silene/genética , Animais , Transtornos do Desenvolvimento Sexual/genética , Flores/anatomia & histologia , Flores/genética , Silene/anatomia & histologia
10.
Plant J ; 49(2): 338-53, 2007 Jan.
Artigo em Inglês | MEDLINE | ID: mdl-17181778

RESUMO

Phytochromes are the primary red/far-red photoreceptors of higher plants, mediating numerous developmental processes throughout the life cycle, from germination to flowering. In seed plants, phytochromes are encoded by a small gene family with each member performing both distinct and redundant roles in mediating physiological responses to light cues. Studies in both eudicot and monocot species have defined a central role for phytochrome B in mediating responses to light in the control of several agronomically important traits, including plant height, transitions to flowering and axillary branch meristem development. Here we characterize Mutator-induced alleles of PhyB1 and a naturally occurring deletion allele of PhyB2 in Zea mays (maize). Using single and double mutants, we show that the highly similar PhyB1 and PhyB2 genes encode proteins with both overlapping and non-redundant functions that control seedling and mature plant traits. PHYB1 and PHYB2 regulate elongation of sheath and stem tissues of mature plants and contribute to the light-mediated regulation of PhyA and Cab gene transcripts. However, PHYB1 and not PHYB2 contributes significantly to the inhibition of mesocotyl elongation under red light, whereas PHYB2 and to a lesser extent PHYB1 mediate the photoperiod-dependent floral transition. This sub functionalization of PHYB activities in maize has probably occurred since the tetraploidization of maize, and may contribute to flowering time variation in modern-day varieties.


Assuntos
Fitocromo B/genética , Proteínas de Plantas/genética , Plântula/genética , Zea mays/genética , Alelos , Northern Blotting , Western Blotting , Elementos de DNA Transponíveis/genética , Flores/genética , Flores/metabolismo , Regulação da Expressão Gênica de Plantas/efeitos da radiação , Luz , Modelos Genéticos , Mutação , Fenótipo , Fitocromo A/genética , Fitocromo A/metabolismo , Fitocromo B/metabolismo , Fitocromo B/fisiologia , Proteínas de Plantas/metabolismo , Proteínas de Plantas/fisiologia , Isoformas de Proteínas/genética , Isoformas de Proteínas/metabolismo , Isoformas de Proteínas/fisiologia , Plântula/metabolismo , Zea mays/metabolismo
11.
Ann Bot ; 95(1): 247-54, 2005 Jan.
Artigo em Inglês | MEDLINE | ID: mdl-15596472

RESUMO

BACKGROUND AND AIMS: The overall goal of this paper is to construct an overview of the genetic basis for flower size evolution in Silene latifolia. It aims to examine the relationship between the molecular bases for flower size and the underlying assumption of quantitative genetics theory that quantitative variation is ultimately due to the impact of a number of structural genes. SCOPE: Previous work is reviewed on the quantitative genetics and potential for response to selection on flower size, and the relationship between flower size and nuclear DNA content in S. latifolia. These earlier findings provide a framework within which to consider more recent analyses of a joint quantitative trait loci (QTL) analysis of flower size and DNA content in this species. KEY RESULTS: Flower size is a character that fits the classical quantitative genetics model of inheritance very nicely. However, an earlier finding that flower size is correlated with nuclear DNA content suggested that quantitative aspects of genome composition rather than allelic substitution at structural loci might play a major role in the evolution of flower size. The present results reported here show that QTL for flower size are correlated with QTL for DNA content, further corroborating an earlier result and providing additional support for the conclusion that localized variations in DNA content underlie evolutionary changes in flower size. CONCLUSIONS: The search image for QTL should be broadened to include overall aspects of genome regulation. As we prepare to enter the much-heralded post-genomic era, we also need to revisit our overall models of the relationship between genotype and phenotype to encompass aspects of genome structure and composition beyond structural genes.


Assuntos
Evolução Biológica , Flores/genética , Genoma de Planta , Silene/genética , Núcleo Celular/genética , Cruzamentos Genéticos , DNA de Plantas/análise , DNA de Plantas/genética , Citometria de Fluxo , Flores/anatomia & histologia , Locos de Características Quantitativas , Seleção Genética
12.
Proc Biol Sci ; 271 Suppl 6: S493-7, 2004 Dec 07.
Artigo em Inglês | MEDLINE | ID: mdl-15801614

RESUMO

Nuclear DNA content variation over orders of magnitude across species has been attributed to 'junk' repetitive DNA with limited adaptive significance. By contrast, our previous work on Silene latifolia showed that DNA content is negatively correlated with flower size, a character of clear adaptive relevance. The present paper explores this relationship in a broader phylogenetic context to investigate the long-term evolutionary impacts of DNA content variation. The relationship between nuclear DNA content and phenotype variation was determined for four closely related species of Silene section Elisanthe (Caryophyllaceae). In addition to a consistent sexual dimorphism in DNA content across all of the species, we found DNA content variation among populations within, as well as among, species. We also found a general trend towards a negative correlation between DNA content and flower and leaf size over all four species, within males and females as well as overall. These results indicate that repetitive DNA may play a role in long-term phenotypic evolution.


Assuntos
Evolução Biológica , DNA Intergênico/genética , Genoma de Planta , Fenótipo , Silene/genética , Análise de Variância , Animais , Composição de Bases , Núcleo Celular/genética , Citometria de Fluxo , Flores/anatomia & histologia , Flores/genética , Portugal , Propídio , Fatores Sexuais , Espanha , Especificidade da Espécie
13.
Plant Physiol ; 133(4): 1578-91, 2003 Dec.
Artigo em Inglês | MEDLINE | ID: mdl-14645729

RESUMO

As an emerging maize (Zea mays) seedling senses light, there is a decrease in the rate of mesocotyl elongation, an induction of root growth, and an expansion of leaves. In leaf tissues, mesophyll and bundle sheath cell fate is determined, and the proplastids of each differentiate into the dimorphic chloroplasts typical of each cell type. Although it has been inferred from recent studies in several model plant species that multiple photoreceptor systems mediate this process, surprisingly little is known of light signal transduction in maize. Here, we examine two photomorphogenic responses in maize: inhibition of mesocotyl elongation and C4 photosynthetic differentiation. Through an extensive survey of white, red, far-red, and blue light responses among a diverse collection of germplasm, including a phytochrome-deficient mutant elm1, we show that light response is a highly variable trait in maize. Although all inbreds examined appear to have a functional phytochrome signal transduction pathway, several lines showed reduced sensitivity to blue light. A significant correlation was observed between light response and subpopulation, suggesting that light responsiveness may be a target of artificial selection. An examination of C4 gene expression patterns under various light regimes in the standard W22 inbred and elm1 indicate that cell-specific patterns of C4 gene expression are maintained in fully differentiated tissues independent of light quality. To our knowledge, these findings represent the first comprehensive survey of light response in maize and are discussed in relation to maize breeding strategies.


Assuntos
Morfogênese/fisiologia , Zea mays/crescimento & desenvolvimento , Sequência de Bases , Diferenciação Celular/efeitos da radiação , Primers do DNA , Regulação da Expressão Gênica de Plantas/genética , Análise dos Mínimos Quadrados , Luz , Fotossíntese/fisiologia , Brotos de Planta/crescimento & desenvolvimento , Brotos de Planta/efeitos da radiação , Sementes , Zea mays/citologia , Zea mays/genética , Zea mays/efeitos da radiação
14.
Plant Physiol ; 130(1): 155-63, 2002 Sep.
Artigo em Inglês | MEDLINE | ID: mdl-12226496

RESUMO

To begin the functional dissection of light signal transduction pathways of maize (Zea mays), we have identified and characterized the light-sensing mutant elm1 (elongated mesocotyl1). Seedlings homozygous for elm1 are pale green, show pronounced elongation of the mesocotyl, and fail to de-etiolate under red or far-red light. Etiolated elm1 mutants contain no spectrally active phytochrome and do not deplete levels of phytochrome A after red-light treatment. High-performance liquid chromatography analyses show that elm1 mutants are unable to convert biliverdin IX alpha to 3Z-phytochromobilin, preventing synthesis of the phytochrome chromophore. Despite the impairment of the phytochrome photoreceptors, elm1 mutants can be grown to maturity in the field. Mature plants retain aspects of the seedling phenotype and flower earlier than wild-type plants under long days. Thus, the elm1 mutant of maize provides the first direct evidence for phytochrome-mediated modulation of flowering time in this agronomically important species.


Assuntos
Biliverdina/análogos & derivados , Fitocromo/metabolismo , Caules de Planta/crescimento & desenvolvimento , Zea mays/crescimento & desenvolvimento , Biliverdina/metabolismo , Carotenoides/metabolismo , Clorofila/metabolismo , Cloroplastos/metabolismo , Escuridão , Luz , Complexos de Proteínas Captadores de Luz , Mutação , Fotossíntese/fisiologia , Fotossíntese/efeitos da radiação , Complexo de Proteínas do Centro de Reação Fotossintética/genética , Complexo de Proteínas do Centro de Reação Fotossintética/metabolismo , Complexo de Proteínas do Centro de Reação Fotossintética/efeitos da radiação , Fitocromo A , Caules de Planta/genética , Caules de Planta/efeitos da radiação , Transdução de Sinais/fisiologia , Zea mays/genética , Zea mays/metabolismo
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