TAC1, a major quantitative trait locus controlling tiller angle in rice

A critical step during rice (Oryza sativa) cultivation is dense planting: a wider tiller angle will increase leaf shade and decrease photosynthesis efficiency, whereas a narrower tiller angle makes for more efficient plant architecture. The molecular basis of tiller angle remains unknown. This research demonstrates that tiller angle is controlled by a major quantitative trait locus, TAC1 (Tiller Angle Control 1). TAC1 was mapped to a 35-kb region on chromosome 9 using a large F(2) population from crosses between an indica rice, IR24, which displays a relatively spread-out plant architecture, and an introgressed line, IL55, derived from japonica rice Asominori, which displays a compact plant architecture with extremely erect tillers. Genetic complementation further identified the TAC1 gene, which harbors three introns in its coding region and a fourth 1.5-kb intron in the 3'-untranslated region. A mutation in the 3'-splicing site of this 1.5-kb intron from 'AGGA' to 'GGGA' decreases the level of tac1, resulting in a compact plant architecture with a tiller angle close to zero. Further sequence verification of the mutation in the 3'-splicing site of the 1.5-kb intron revealed that the tac1 mutation 'GGGA' was present in 88 compact japonica rice accessions and TAC1 with 'AGGA' was present in 21 wild rice accessions and 43 indica rice accessions, all with the spread-out form, indicating that tac1 had been extensively utilized in densely planted rice grown in high-latitude temperate areas and at high altitudes where japonica rice varieties are widely cultivated.     

A Novel Tiller Angle Gene,TAC3, together with TAC1 and D2 Largely Determine the Natural Variation of Tiller Angle in Rice Cultivars

Tiller angle is one of the most important components of the ideal plant architecture that can greatly enhance rice grain yield. Understanding the genetic basis of tiller angle and mining favorable alleles will be helpful for breeding new plant-type varieties. Here, we performed genome-wide association studies (GWAS) to identify genes controlling tiller angle using 529 diverse accessions of Oryza sativa including 295 indica and 156 japonica accessions in two environments. We identified 7 common quantitative trait loci (QTLs), including the previously reported major gene Tiller Angle Control 1 (TAC1), in the two environments, 10 and 13 unique QTLs in Hainan and Wuhan, respectively. More QTLs were identified in indica than in japonica, and three major QTLs (qTA3, qTA1b/DWARF2 (D2) and qTA9c/TAC1) were fixed in japonica but segregating in indica, which explained the wider variation observed in indica compared with that in japonica. No common QTLs were identified between the indica and japonica subpopulations. Mutant analysis for the candidate gene of qTA3 on chromosome 3 indicated a novel gene, Tiller Angle Control 3 (TAC3), encoding a conserved hypothetical protein controlling tiller angle. TAC3 is preferentially expressed in the tiller base. The ebisu dwarf (d2) mutant exhibited a decreased tiller angle, in addition to its previously described abnormal phenotype. A nucleotide diversity analysis revealed that TAC3, D2 and TAC1 have been subjected to selection during japonica domestication. A haplotype analysis identified favorable alleles of TAC3, D2 and TAC1, which may be used for breeding plants with an ideal architecture. In conclusion, there is a diverse genetic basis for tiller angle between the two subpopulations, and it is the novel gene TAC3 together with TAC1, D2, and other newly identified genes in this study that controls tiller angle in rice cultivars.     

Natural variation in Tiller Number 1 affects its interaction with TIF1 to regulate tillering in rice

Tiller number per plant—a cardinal component of ideal plant architecture—affects grain yield potential. Thus, alleles positively affecting tillering must be mined to promote genetic improvement. Here, we report a Tiller Number 1 (TN1) protein harbouring a bromo-adjacent homology domain and RNA recognition motifs, identified through genome-wide association study of tiller numbers. Natural variation in TN1 affects its interaction with TIF1 (TN1 interaction factor 1) to affect DWARF14 expression and negatively regulate tiller number in rice. Further analysis of variations in TN1 among indica genotypes according to geographical distribution revealed that low-tillering varieties with TN1-hap^L are concentrated in Southeast Asia and East Asia, whereas high-tillering varieties with TN1-hap^H are concentrated in South Asia. Taken together, these results indicate that TN1 is a tillering regulatory factor whose alleles present apparent preferential utilization across geographical regions. Our findings advance the molecular understanding of tiller development.     

Molecular Evolution of the TAC1 Gene from Rice （Oryza sativa L.）

Tiller angle is a key feature of the architecture of cultivated rice(Oryza sativa),since it determines planting density and influences rice yield.Our previous work identified Tiller Angle Control 1(TACl) as a major quantitative trait locus that controls rice tiller angle.To further clarify the evolutionary characterization of the TACl gene,we compared a TACl-containing 3164-bp genomic region among 113 cultivated varieties and 48 accessions of wild rice,including 43 accessions of O.rufipogon and five accessions of O.nivara.Only one single nucleotide polymorphism(SNP),a synonymous substitution,was detected in TACl coding regions of the cultivated rice varieties, whereas one synonymous and one nonsynonymous SNP were detected among the TACl coding regions of wild rice accessions.These data indicate that little natural mutation and modification in the TACl coding region occurred within the cultivated rice and its progenitor during evolution.Nucleotide diversities in the TACl gene regions of O.sativa and O.rufipogon of 0.00116 and 0.00112,respectively, further indicate that TACl has been highly conserved during the course of rice domestication.A functional nucleotide polymorphism (FNP) of TACl was only found in the japonica rice group.A neutrality test revealed strong selection,especially in the 3’-flanking region of the TACl coding region containing the FNP in the japonica rice group.However,no selection occurred in the indica and wild-rice groups.A phylogenetic tree derived from TACl sequence analysis suggests that the indica and japonica subspecies arose independently during the domestication of wild rice.     

Understanding the Regulatory Mechanisms of Rice Tiller Angle,Then and Now

Rice is one of the most important crops worldwide, whose yield is vital to human nutrition in the context of a rapidly growing world population. Plant architecture significantly affects grain yield, which is to a large extent determined by tiller angle and tiller number. Tiller angle is the angle between the primary tiller and the main culm. Its regulation is complex and is influenced by multiple environmental and genetic factors. This review provides an overview of the regulation of tiller angle in rice, with particular focus on the roles of the growth environment and method of cultivation; phytohormones such as auxin, gibberellins, and strigolactones; gravity; and genes related to the control of tiller angle. The major research foci and the outlook for research into the regulation of tiller angle in rice are discussed.

     

PpeTAC1 promotes the horizontal growth of branches in peach trees and is a member of a functionally conserved gene family found in diverse plants species

Trees are capable of tremendous architectural plasticity, allowing them to maximize their light exposure under highly competitive environments. One key component of tree architecture is the branch angle, yet little is known about the molecular basis for the spatial patterning of branches in trees. Here, we report the identification of a candidate gene for the br mutation in Prunus persica (peach) associated with vertically oriented growth of branches, referred to as ‘pillar’ or ‘broomy’. Ppa010082, annotated as hypothetical protein in the peach genome sequence, was identified as a candidate gene for br using a next generation sequence‐based mapping approach. Sequence similarity searches identified rice TAC1 (tiller angle control 1) as a putative ortholog, and we thus named it PpeTAC1. In monocots, TAC1 is known to lead to less compact growth by increasing the tiller angle. In Arabidopsis, an attac1 mutant showed more vertical branch growth angles, suggesting that the gene functions universally to promote the horizontal growth of branches. TAC1 genes belong to a gene family (here named IGT for a shared conserved motif) found in all plant genomes, consisting of two clades: one containing TAC1‐like genes; the other containing LAZY1, which contains an EAR motif, and promotes vertical shoot growth in Oryza sativa (rice) and Arabidopsis through influencing polar auxin transport. The data suggest that IGT genes are ancient, and play conserved roles in determining shoot growth angles in plants. Understanding how IGT genes modulate branch angles will provide insights into how different architectural growth habits evolved in terrestrial plants.     

LAZY1 controls rice shoot gravitropism through regulating polar auxin transport

Tiller angle of rice （Oryza sativa L.） is an important agronomic trait that contributes to grain production, and has long attracted attentions of breeders for achieving ideal plant architecture to improve grain yield. Although enormous efforts have been made over the past decades to study mutants with extremely spreading or compact tillers, the molecular mechanism underlying the control of tiller angle of cereal crops remains unknown. Here we report the cloning of the LAZY1 （LA1） gene that regulates shoot gravitropism by which the rice tiller angle is controlled. We show that LA1, a novel grass-specific gene, is temporally and spatially expressed, and plays a negative role in polar auxin transport （PAT）. Loss-of-function of LA1 enhances PAT greatly and thus alters the endogenous IAA distribution in shoots, leading to the reduced gravitropism, and therefore the tiller-spreading phenotype of rice plants.     

Natural variation and genetic analysis of the tiller angle gene MsTAC1 in Miscanthus sinensis

Biomass yield is an important target trait in Miscanthus breeding for desirable energy crops. Tiller angle is a key trait of plant architecture because it determines planting density and further influences biomass yield through affecting photosynthesis efficiency. TAC1, a major gene involved in tiller and leaf angle control in rice and maize, respectively, has been extensively studied. Nucleotide variation at this gene, MsTAC1, was investigated in 33 Miscanthus sinensis accessions collected from different areas in China, and one genotype of Miscanthus × giganteus. A total of 136 loci, including 129 single base substitutions and seven InDels, occurred within the MsTAC1 gene of 1,874 bp. The genetic diversity at MsTAC1 is characterized by high nucleotide diversity (π value) and high heterozygosity. Clustering analysis indicated that the phylogenetic tree of the 33 M. sinensis accessions was correlated with their geographical sites of origin. The neutrality test revealed no strong selection pressure on coding and non-coding region variations of the MsTAC1 gene in the accessions. Phenotype evaluations were conducted for tiller angle and five other traits in the Miscanthus panels in the first two growth years of 2009 and 2010. Analysis of variance showed significant phenotypic variations in the examined traits. Association analysis using 246 markers detected 88 loci associated with all the test traits in either 1 or 2 years, and 11 of the 88 were year reproducible and thus reliable. These associations indicate that the variation of MsTAC1 affects the phenotypic value of the tiller angle, tiller number and biomass yield, suggesting that allelic variation in MsTAC1 affects multiple traits and demonstrates its significance in Miscanthus breeding programs.     

OsmiR167a‐targeted auxin response factors modulate tiller angle via fine‐tuning auxin distribution in rice

Rice tiller angle determines plant growth density and further contributes grain production. Although a few genes have been characterized to regulate tiller angle in rice, the molecular mechanism underlying the control of tiller angle via microRNA is poorly understood. Here, we report that rice tiller angle is controlled by OsmiR167a‐targeted auxin response factors OsARF12, OsARF17 and OsARF25. In the overexpression of OsMIR167a plants, the expression of OsARF12, OsARF17 and OsARF25 was severely repressed and displayed larger tiller angle as well as the osarf12/osarf17 and osarf12/ osarf25 plants. In addition, those plants showed compromised abnormal auxin distribution and less sensitive to gravity. We also demonstrate that OsARF12, OsARF17 and OsARF25 function redundantly and might be involved in HSFA2D and LAZY1‐dependent asymmetric auxin distribution pathway to control rice tiller angle. Our results reveal that OsmiR167a represses its targets, OsARF12, OsARF17 and OsARF25, to control rice tiller angle by fine‐tuning auxin asymmetric distribution in shoots.     

OsPIN5b modulates rice (Oryza sativa) plant architecture and yield by changing auxin homeostasis,transport and distribution

Plant architecture attributes such as tillering, plant height and panicle size are important agronomic traits that determine rice (Oryza sativa) productivity. Here, we report that altered auxin content, transport and distribution affect these traits, and hence rice yield. Overexpression of the auxin efflux carrier‐like gene OsPIN5b causes pleiotropic effects, mainly reducing plant height, leaf and tiller number, shoot and root biomass, seed‐setting rate, panicle length and yield parameters. Conversely, reduced expression of OsPIN5b results in higher tiller number, more vigorous root system, longer panicles and increased yield. We show that OsPIN5b is an endoplasmic reticulum (ER) ‐localized protein that participates in auxin homeostasis, transport and distribution in vivo. This work describes an example of an auxin‐related gene where modulating its expression can simultaneously improve plant architecture and yield potential in rice, and reveals an important effect of hormonal signaling on these traits.     

植物重力反应的分子调控机制

重力是调节植物生长发育和形态建成的重要环境因子。植物感受到重力刺激后可以通过重力反应来协调自身各个器官的生长方向与重力方向之间的最适角度。植物重力反应过程分为重力信号的感受、重力信号的转导、生长素不对称分布的形成和重力反应器官的弯曲生长4个阶段。近年来,随着大量重力反应缺陷突变体的鉴定及其控制基因的功能解析,重力信号的感受和生长素不对称分布的分子机制等方面的研究取得了重要进展。作为植物适应环境变化的重要手段之一,重力反应还可以通过调节水稻(Oryza sativa L.)的分蘖角度实现对水稻株型和产量的调控。因此,研究植物的重力反应,不仅有助于解析植物生长发育的调控机制,对于作物株型的改良也具有重要的指导意义。然而,重力反应的分子机制及其调控网络仍不清楚。本文综述了近年来植物重力反应的调控机理及其调控水稻分蘖角度的作用机制,并对该领域未来的研究方向和热点进行了展望。     

Demographic variability in tiller populations of two perennial pampa grasses

Abstract. The general objectives of this study were: (1) to investigate the importance of internal influences in regulating the tiller dynamics in natural populations of the warm-season perennial grasses Paspalum dilatatum and Sporobolus indicus, coexisting in Argentine flooded pampa, in as much as they act independently of the underlying external environment, and (2) to evaluate the extent to which interactions between internal and external factors affect the variation in tiller dynamics within such populations. Within-population variation in seasonal development of plants and tillers with different neighbour composition was studied for an annual growth cycle. Tiller survival and tillering were significantly influenced by tiller size. Tiller age influenced tiller fate, as suggested by the additive effects of age and size of tillers. These relationships varied with season and with species. Size and age of tillers showed additive effects with their neighbouring species on the tiller fate of P. dilatatum, but the effects of age and size of S. indicus changed according their neighbourhood. Tiller survival of S. indicus during the early growth season was more size-dependent when the cold-season species Poa lanigera, was the principal neighbour. Flowering and tillering probabilities were positively related through their common positive responses to tiller size. Tiller survival and recruitment between different seasons were strongly related. Independently of neighbour composition, tiller survival was generally inversely related to tiller recruitment in previous seasons. Therefore, significant density-dependent mortality of tillers was found for both species during the early summer when tiller density was expressed by basal area units.     

LAZY3 interacts with LAZY2 to regulate tiller angle by modulating shoot gravity perception in rice

Starch biosynthesis in gravity-sensing tissues of rice shoot determines the magnitude of rice shoot gravitropism and thus tiller angle. However, the molecular mechanism underlying starch biosynthesis in rice gravity-sensing tissues is still unclear. We characterized a novel tiller angle gene LAZY3 (LA3) in rice through map-based cloning. Biochemical, molecular and genetic studies further demonstrated the essential roles of LA3 in gravity perception of rice shoot and tiller angle control. The shoot gravitropism and lateral auxin transport were defective in la3 mutant upon gravistimulation. We showed that LA3 encodes a chloroplast-localized tryptophan-rich protein associated with starch granules via Tryptophan-rich region (TRR) domain. Moreover, LA3 could interact with the starch biosynthesis regulator LA2, determining starch granule formation in shoot gravity-sensing tissues. LA3 and LA2 negatively regulate tiller angle in the same pathway acting upstream of LA1 to mediate asymmetric distribution of auxin. Our study defined LA3 as an indispensable factor of starch biosynthesis in rice gravity-sensing tissues that greatly broadens current understanding in the molecular mechanisms underlying the starch granule formation in gravity-sensing tissues, and provides new insights into the regulatory mechanism of shoot gravitropism and rice tiller angle.     

Ectopic expression of foxtail millet zip-like gene, <Emphasis Type="Italic">SiPf40</Emphasis>, in transgenic rice plants causes a pleiotropic phenotype affecting tillering,vascular distribution and root development

Plant architecture determines grain production in rice (Oryza sativa) and is affected by important agronomic traits such as tillering, plant height, and panicle morphology. Many key genes involved in controlling the initiation and outgrowth of axillary buds, the elongation of stems, and the architecture of inflorescences have been isolated and analyzed. Previous studies have shown that SiPf40, which was identified from a foxtail millet (Setaria italica) immature seed cDNA library, causes extra branches and tillers in SiPf40-transgenic tobacco and foxtail millet, respectively. To reconfirm its function, we generated transgenic rice plants overexpressing SiPf40 under the control of the ubiquitin promoter. SiPf40-overexpressing transgenic plants have a greater tillering number and a wider tiller angle than wild-type plants. Their root architecture is modified by the promotion of lateral root development, and the distribution of xylem and phloem in the vascular bundle is affected. Analysis of hormone levels showed that the ratios of indole-3-acetic acid/zeatin (IAA/ZR) and IAA/gibberellic acid (IAA/GA) decreased in SiPf40-transgenic plants compared with wild-type plants. These findings strongly suggest that SiPf40 plays an important role in plant architecture.     

Growth and carbon allocation of Agropyron desertorum following autumn defoliation

Summary Growth and carbon allocation of a cool season tussock grass, Agropyron desertorum, following defoliation of newly initiated tillers in the autumn of 1988 and 1989 were investigated. Tiller density and mortality, reproductive shoot density, root density, biomass, individual tiller weight, carbon allocation, and soil water depletion were used to evaluate the response of A. desertorum to autumn grazing. Tiller recruitment was lower in the autumn-defoliated treatment in both years compared with the control because of the cessation of tiller development following autumn defoliation. Autumn defoliation also significantly reduced the movement of ¹³C to the roots in 1988 but not in 1989. Soils were cooler and drier in 1989. Other plant growth measurements and soil water depletion rates were not different between treatments. Autumn defoliation in 1988 did not influence tiller recruitment in the following autumn. Two consecutive years of autumn defoliation did not affect tiller overwinter mortality or peak standing crop in 1990.     

Characterization of a novel high-tillering dwarf 3 mutant in rice

Tiller number and culm length are important components of plant architecture and determinate grain production in rice.A line SIL046,derived from an introgression lines population developed by an accession of common wild rice (Oryza rufipogon Griff.) and a high-yielding indica cultivar Guichao 2 (Oryza sativa L.).exhibits a higher tiller number and shorter culm length phenotype than the recipient parent Guichao 2 (GC2).Genetic analysis showed that the high-tillering dwarf phenotype was controlled by a novel single recessive gene,referred to as the high-tillering dwarf3 (htd3),which located within the genetic distance of 13.4 cM between SSR makers RM7003 and RM277 on chromosome 12.By means of fine-mapping strategy,we mapped HTD3 gene within the genetic distance of 2.5 cM and the physical distance of 3100 kb in the centromere of chromosome 12.Further identification of HTD3 gene would provide a new opportunity to uncover the molecular mechanism of the development of culm and tiller,two important components of yields in rice.