RESUMEN
Seed vigor in crops is important in terms of improving grain quality and germplasm conservation; however, little is known about its regulatory mechanisms through the encoded proteome and gene network. Comparative analyses of transcriptome (RNA sequencing [RNA-seq]) and broadly targeted metabolic profiling of two subspecific rice cultivars with distinct seed vigor during accelerated aging revealed various biological pathways and metabolic processes as key influences explaining trait differences. RNA-seq coexpression regulatory network analyses identified several transcription factors, including bZIP23 and bZIP42, that act as nodes in the gene network. Importantly, transgenic seeds of overexpression of bZIP23 enhanced seed vigor, whereas its gene knockout reduced seed vigor, suggesting that the protein it encodes functions as a positive regulator. Similarly, overexpression and knockout of PER1A that encodes a key player in the detoxification pathway enhanced and decreased seed vigor, respectively. We further demonstrated a direct interaction of the PER1A promoter with bZIP23 in seeds, which activates the expression of PER1A, and the genetic evidence suggested that bZIP23 most likely functions in a common pathway with and acts upstream of PER1A to modulate seed vigor. In addition, the control of seed vigor by the bZIP23-PER1A module was connected with that of the abscisic acid signaling pathway. Collectively, we revealed the genetic architecture of variation in seed vigor and uncovered the bZIP23-PER1A-mediated detoxification pathway that enhances the trait in rice.
Asunto(s)
Genoma de Planta , Vigor Híbrido , Metaboloma , Oryza/embriología , Peroxirredoxinas/metabolismo , Proteínas de Plantas/metabolismo , Semillas/fisiología , Ácido Abscísico/metabolismo , Perfilación de la Expresión Génica , Regulación de la Expresión Génica de las Plantas , Genes de Plantas , Oryza/genética , Oryza/metabolismo , Semillas/metabolismo , Transducción de SeñalRESUMEN
From its tropical origin in southwestern Mexico, maize spread over a wide latitudinal cline in the Americas. This feat defies the rule that crops are inhibited from spreading easily across latitudes. How the widespread latitudinal adaptation of maize was accomplished is largely unknown. Through positional cloning and association mapping, we resolved a flowering-time quantitative trait locus to a Harbinger-like transposable element positioned 57 kb upstream of a CCT transcription factor (ZmCCT9). The Harbinger-like element acts in cis to repress ZmCCT9 expression to promote flowering under long days. Knockout of ZmCCT9 by CRISPR/Cas9 causes early flowering under long days. ZmCCT9 is diurnally regulated and negatively regulates the expression of the florigen ZCN8, thereby resulting in late flowering under long days. Population genetics analyses revealed that the Harbinger-like transposon insertion at ZmCCT9 and the CACTA-like transposon insertion at another CCT paralog, ZmCCT10, arose sequentially following domestication and were targeted by selection for maize adaptation to higher latitudes. Our findings help explain how the dynamic maize genome with abundant transposon activity enabled maize to adapt over 90° of latitude during the pre-Columbian era.
Asunto(s)
Adaptación Fisiológica/genética , Regulación de la Expresión Génica de las Plantas/fisiología , Proteínas de Plantas/metabolismo , Factores de Transcripción/metabolismo , Zea mays/genética , Zea mays/fisiología , Clonación Molecular , Flores/genética , Flores/fisiología , Eliminación de Gen , Genoma de Planta , Proteínas de Plantas/genéticaRESUMEN
Flowering time is a major determinant of the local adaptation of plants. Although numerous loci affecting flowering time have been mapped in maize, their underlying molecular mechanisms and roles in adaptation remain largely unknown. Here, we report the identification and characterization of MADS-box transcription factor ZmMADS69 that functions as a flowering activator through the ZmRap2.7-ZCN8 regulatory module and contributes to adaptation. We show that ZmMADS69 underlies a quantitative trait locus controlling the difference in flowering time between maize and its wild ancestor, teosinte. Maize ZmMADS69 allele is expressed at a higher level at floral transition and confers earlier flowering than the teosinte allele under long days and short days. Overexpression of ZmMADS69 causes early flowering, while a transposon insertion mutant of ZmMADS69 exhibits delayed flowering. ZmMADS69 shows pleiotropic effects for multiple traits of agronomic importance. ZmMADS69 functions upstream of the flowering repressor ZmRap2.7 to downregulate its expression, thereby relieving the repression of the florigen gene ZCN8 and causing early flowering. Population genetic analyses showed that ZmMADS69 was a target of selection and may have played an important role as maize spread from the tropics to temperate zones. Our findings provide important insights into the regulation and adaptation of flowering time.
Asunto(s)
Flores/fisiología , Proteínas de Dominio MADS/metabolismo , Proteínas de Plantas/metabolismo , Zea mays/metabolismo , Adaptación Fisiológica/genética , Mapeo Cromosómico , Florigena , Regulación de la Expresión Génica de las Plantas , Técnicas de Inactivación de Genes , Genética de Población , Proteínas de Dominio MADS/genética , Proteínas de Plantas/genética , Plantas Modificadas Genéticamente , Sitios de Carácter Cuantitativo , Selección Genética , Zea mays/genéticaRESUMEN
Maize (Zea mays) tassels underwent profound morphological changes during maize domestication and improvement. Although a number of genes affecting maize inflorescence development have been identified, the genetic basis of the morphological changes in maize tassels since domestication is not well understood. Here, using a large population of 866 maize-teosinte BC2 S3 recombinant inbred lines genotyped using 19 838 single nucleotide polymorphism (SNP) markers, we performed high-resolution quantitative trait locus (QTL) mapping for five tassel morphological traits. We showed that the five tassel traits were associated with different genetic architecture features. Known genes for maize inflorescence development identified by mutagenesis were significantly enriched in the tassel trait QTLs, and many of these genes, including ramosa1 (ra1), barren inflorescence2 (bif2), unbranched2 (ub2), zea floricaula leafy2 (zfl2) and barren stalk fastigiate1 (baf1), showed evidence of selection. An in-depth nucleotide diversity analysis at the bif2 locus identified strong selection signatures in the 5'-regulatory region. We also found that several known flowering time genes co-localized with tassel trait QTLs. A further association analysis indicated that the maize photoperiod gene ZmCCT was significantly associated with tassel size variation. Using near-isogenic lines, we narrowed down a major-effect QTL for tassel length, qTL9-1, to a 513-kb physical region. These results provide important insights into the genetic architecture that controls maize tassel evolution.
Asunto(s)
Domesticación , Flores/anatomía & histología , Zea mays/anatomía & histología , Zea mays/genética , Flores/genética , Flores/fisiología , Genes de Plantas , Endogamia , Inflorescencia/genética , Inflorescencia/crecimiento & desarrollo , Fenotipo , Mapeo Físico de Cromosoma , Sitios de Carácter Cuantitativo/genética , Carácter Cuantitativo Heredable , Recombinación Genética/genética , Selección Genética , Factores de TiempoRESUMEN
The number of leaves and their distributions on plants are critical factors determining plant architecture in maize (Zea mays), and leaf number is frequently used as a measure of flowering time, a trait that is key to local environmental adaptation. Here, using a large set of 866 maize-teosinte BC2 S3 recombinant inbred lines genotyped by using 19,838 single nucleotide polymorphism markers, we conducted a comprehensive genetic dissection to assess the genetic architecture of leaf number and its genetic relationship to flowering time. We demonstrated that the two components of total leaf number, the number of leaves above (LA) and below (LB) the primary ear, were under relatively independent genetic control and might be subject to differential directional selection during maize domestication and improvement. Furthermore, we revealed that flowering time and leaf number are commonly regulated at a moderate level. The pleiotropy of the genes ZCN8, dlf1 and ZmCCT on leaf number and flowering time were validated by near-isogenic line analysis. Through fine mapping, qLA1-1, a major-effect locus that specifically affects LA, was delimited to a region with severe recombination suppression derived from teosinte. This study provides important insights into the genetic basis of traits affecting plant architecture and adaptation. The genetic independence of LA from LB enables the optimization of leaf number for ideal plant architecture breeding in maize.
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Flores/fisiología , Hojas de la Planta/anatomía & histología , Zea mays/genética , Zea mays/fisiología , Cruzamientos Genéticos , Flores/genética , Estudios de Asociación Genética , Endogamia , Fenotipo , Mapeo Físico de Cromosoma , Hojas de la Planta/genética , Sitios de Carácter Cuantitativo/genética , Reproducibilidad de los Resultados , Factores de TiempoRESUMEN
Studies that investigated the genetic basis of source and sink related traits have been widely conducted. However, the vascular system that links source and sink received much less attention. When maize was domesticated from its wild ancestor, teosinte, the external morphology has changed dramatically; however, less is known for the internal anatomy changes. In this study, using a large maize-teosinte experimental population, we performed a high-resolution quantitative trait locus (QTL) mapping for the number of vascular bundle in the uppermost internode of maize stem. The results showed that vascular bundle number is dominated by a large number of small-effect QTLs, in which a total of 16 QTLs that jointly accounts for 52.2% of phenotypic variation were detected, with no single QTL explaining more than 6% of variation. Different from QTLs for typical domestication traits, QTLs for vascular bundle number might not be under directional selection following domestication. Using Near Isogenic Lines (NILs) developed from heterogeneous inbred family (HIF), we further validated the effect of one QTL qVb9-2 on chromosome 9 and fine mapped the QTL to a 1.8-Mb physical region. This study provides important insights for the genetic architecture of vascular bundle number in maize stem and sets basis for cloning of qVb9-2.
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Mapeo Cromosómico/métodos , Tallos de la Planta/genética , Haz Vascular de Plantas/anatomía & histología , Haz Vascular de Plantas/genética , Sitios de Carácter Cuantitativo/genética , Zea mays/genética , Emparejamiento Base/genética , Estudios de Asociación Genética , Genoma de Planta , Endogamia , Escala de Lod , Fenotipo , Recombinación Genética/genéticaRESUMEN
Maxillary sinus floor augmentation using lateral window and crestal technique is considered as predictable methods to increase the residual bone height; however, this surgery is commonly complicated by Schneiderian membrane perforation, which is closely related to anatomical factors. This article aimed to assess anatomical factors on successful augmentation procedures. After review of the current evidence on sinus augmentation techniques, anatomical factors related to the stretching potential of Schneiderian membrane were assessed and a decision tree for the rational choice of surgical approaches was proposed. Schneiderian membrane perforation might occur when local tension exceeds its stretching potential, which is closely related to anatomical variations of the maxillary sinus. Choice of a surgical approach and clinical outcomes are influenced by the stretching potential of Schneiderian membrane. In addition to the residual bone height, clinicians should also consider the stretching potential affected by the membrane health condition, the contours of the maxillary sinus, and the presence of antral septa when evaluating the choice of surgical approaches and clinical outcomes.
Asunto(s)
Elevación del Piso del Seno Maxilar , Árboles de DecisiónRESUMEN
Input-Output (IO) data describing supply-demand relationships between buyers and sellers for goods and services within an economy have been used not only in economics but also in scientific, environmental, and interdisciplinary research. However, most conventional IO data are highly aggregated, resulting in challenges for researchers and practitioners who face complex issues in large countries such as China, where firms within the same IO sector may have significant differences in technologies across subnational regions and different ownerships. This paper is the first attempt to compile China's interprovincial IO (IPIO) tables with separate information for mainland China-, Hong Kong, Macau, Taiwan-, and foreign-owned firms inside each province/industry pair. To do this, we collect relevant Chinese economic census data, firm surveys, product level Custom trade statistics, and firm value-added tax invoices and consistently integrate them into a 42-sector, 31-province IO account covering 5 benchmark years between 1997-2017. This work provides a solid foundation for a diverse range of innovative IO-based research in which firm heterogeneity information about location and ownership matters.
RESUMEN
Gene expression regulation plays an important role in controlling plant phenotypes and adaptation. Here, we report a comprehensive assessment of gene expression variation through the transcriptome analyses of a large maize-teosinte experimental population. Genome-wide mapping identified 25 660 expression quantitative trait loci (eQTL) for 17 311 genes, capturing an unprecedented range of expression variation. We found that local eQTL were more frequently mapped to adjacent genes, displaying a mode of expression piggybacking, which consequently created co-regulated gene clusters. Genes within the co-regulated gene clusters tend to have relevant functions and shared chromatin modifications. Distant eQTL formed 125 significant distant eQTL hotspots with their targets significantly enriched in specific functional categories. By integrating different sources of information, we identified putative trans- regulators for a variety of metabolic pathways. We demonstrated that the bHLH transcription factor R1 and hexokinase HEX9 might act as crucial regulators for flavonoid biosynthesis and glycolysis, respectively. Moreover, we showed that domestication or improvement has significantly affected global gene expression, with many genes targeted by selection. Of particular interest, the Bx genes for benzoxazinoid biosynthesis may have undergone coordinated cis-regulatory divergence between maize and teosinte, and a transposon insertion that inactivates Bx12 was under strong selection as maize spread into temperate environments with a distinct herbivore community.