Domestication rewired gene expression and nucleotide diversity patterns in tomato

Plant domestication has led to considerable phenotypic modifications from wild species to modern varieties. However, although changes in key traits have been well documented, less is known about the underlying molecular mechanisms, such as the reduction of molecular diversity or global gene co‐expression patterns. In this study, we used a combination of gene expression and population genetics in wild and crop tomato to decipher the footprints of domestication. We found a set of 1729 differentially expressed genes (DEG) between the two genetic groups, belonging to 17 clusters of co‐expressed DEG, suggesting that domestication affected not only individual genes but also regulatory networks. Five co‐expression clusters were enriched in functional terms involving carbohydrate metabolism or epigenetic regulation of gene expression. We detected differences in nucleotide diversity between the crop and wild groups specific to DEG. Our study provides an extensive profiling of the rewiring of gene co‐expression induced by the domestication syndrome in one of the main crop species.     

Genomic screening for artificial selection during domestication and improvement in maize

BACKGROUND: Artificial selection results in phenotypic evolution. Maize (Zea mays L. ssp. mays) was domesticated from its wild progenitor teosinte (Zea mays subspecies parviglumis) through a single domestication event in southern Mexico between 6000 and 9000 years ago. This domestication event resulted in the original maize landrace varieties. The landraces provided the genetic material for modern plant breeders to select improved varieties and inbred lines by enhancing traits controlling agricultural productivity and performance. Artificial selection during domestication and crop improvement involved selection of specific alleles at genes controlling key morphological and agronomic traits, resulting in reduced genetic diversity relative to unselected genes. SCOPE: This review is a summary of research on the identification and characterization by population genetics approaches of genes affected by artificial selection in maize. CONCLUSIONS: Analysis of DNA sequence diversity at a large number of genes in a sample of teosintes and maize inbred lines indicated that approx. 2 % of maize genes exhibit evidence of artificial selection. The remaining genes give evidence of a population bottleneck associated with domestication and crop improvement. In a second study to efficiently identify selected genes, the genes with zero sequence diversity in maize inbreds were chosen as potential targets of selection and sequenced in diverse maize landraces and teosintes, resulting in about half of candidate genes exhibiting evidence for artificial selection. Extended gene sequencing demonstrated a low false-positive rate in the approach. The selected genes have functions consistent with agronomic selection for plant growth, nutritional quality and maturity. Large-scale screening for artificial selection allows identification of genes of potential agronomic importance even when gene function and the phenotype of interest are unknown. These approaches should also be applicable to other domesticated species if specific demographic conditions during domestication exist.     

亚洲栽培稻主要驯化性状研究进展

水稻是研究谷类作物驯化的良好材料, 其中种子落粒性消失、休眠性减弱和株型上的变化是水稻驯化过程中的3个关键事件, 造就了高产、发芽整齐及可密植的现代水稻。落粒性丧失一直被认为是野生稻驯化形态学上的最直接证据, 而控制落粒的主要基因Sh4和qSH1分别暗示不同的水稻驯化历史。种子休眠性的减弱适应了现代农业生产上同步发芽的需求, Sdr4、qSD7-1和qSD12基因是目前已知的调控种子休眠性的3个关键位点。野生稻匍匐生长等特点与其长期所在的易变生境有关, 而栽培稻的直立生长形态则适应了农业上密植生产的需要, 受PROG1等基因控制。野生稻的异交特性促进了驯化基因在群体间传播, 而自花授粉则使驯化基因得以稳定遗传, 从而加快人工选择的累积。目前的水稻驯化研究侧重于单基因或一些中性标记, 而对控制驯化性状的网络化通路的进化研究却相对缺乏。随着功能基因组研究的深入, 驯化性状的分子机理将会被全面揭示, 而基于此的网络化通路研究必将更加真实地反应水稻驯化过程。文章综述了水稻关键驯化性状分子机理的研究进展, 为驯化基因网络的研究提供参考, 也为水稻分子设计改良提供新的思路。     

What Has Natural Variation Taught Us about Plant Development,Physiology, and Adaptation?

Nearly 100 genes and functional polymorphisms underlying natural variation in plant development and physiology have been identified. In crop plants, these include genes involved in domestication traits, such as those related to plant architecture, fruit and seed structure and morphology, as well as yield and quality traits improved by subsequent crop breeding. In wild plants, comparable traits have been dissected mainly in Arabidopsis thaliana. In this review, we discuss the major contributions of the analysis of natural variation to our understanding of plant development and physiology, focusing in particular on the timing of germination and flowering, plant growth and morphology, primary metabolism, and mineral accumulation. Overall, functional polymorphisms appear in all types of genes and gene regions, and they may have multiple mutational causes. However, understanding this diversity in relation to adaptation and environmental variation is a challenge for which tools are now available.     

作物驯化的遗传学研究及其在大豆育种中的应用

驯化遗传学是进化生物学研究的一个重要分支, 对作物驯化历程进行深入探索, 可以更好地认识人类早期农耕文明的产生和发展, 了解物种的形成乃至生命的起源和进化, 对指导人们利用野生资源来改良现有的栽培作物和培育新的有价值的品种也具有重要的意义。本文综述了作物驯化的遗传学研究内容和进展, 以及重要作物大豆驯化起源的研究概况, 探讨了目前作物驯化研究对大豆驯化遗传学及野生大豆资源综合利用的指导意义。     

作物驯化的遗传学研究及其在大豆育种中的应用

驯化遗传学是进化生物学研究的一个重要分支,对作物驯化历程进行深入探索,可以更好地认识人类早期农耕文明的产生和发展,了解物种的形成乃至生命的起源和进化,对指导人们利用野生资源来改良现有的栽培作物和培育新的有价值的品种也具有重要的意义。本文综述了作物驯化的遗传学研究内容和进展,以及重要作物大豆驯化起源的研究概况,探讨了目前作物驯化研究对大豆驯化遗传学及野生大豆资源综合利用的指导意义。     

种子胎萌机制研究进展

种子胎萌是内在的遗传基础和外部环境共同作用的结果,受许多基因的调控和植物激素的影响。近些年来,随着分子生物学的快速发展,种子胎萌研究已经深入到分子水平。分子生物学技术的运用,特别是基因的克隆与表达、植物激素的生物合成与信号转导和分子遗传学等手段已成为研究种子胎萌的新工具和新方向。现从种皮色泽基因R、矮杆基因Rht3以及Viviparous(Vp)基因家族等方面就种子胎萌相关基因与胎萌关系进行了综述;并对植物激素脱落酸(ABA)和赤霉素(GA)的生物合成或信号转导在种子胎萌的调控中的作用等方面进行综述。     

泛素-蛋白酶体系统影响植物农艺性状的研究进展

当前人口激增、环境污染、生态破坏等问题层出不穷。改良农艺性状,如提高产量、增强逆境耐受能力,是作物遗传改良的重要目标,是推动农业高质量发展的基础。泛素-蛋白酶体系统（UPS）是一种快速的选择性水解植物体产生的冗余蛋白和被损坏蛋白的体系。目前,研究发现泛素-蛋白酶系统影响植物的发育、生殖和重要的农艺性状：如应对环境胁迫、开花诱导和种子大小等。综述了近年来的研究成果,阐述了蛋白酶体UPS组分和各亚基的重要功能,并描述了泛素-蛋白酶系统是如何影响植物农艺性状的。最后,讨论了未来的研究热点和利用UPS改良作物的潜力。     

Selection on domestication traits and quantitative trait loci in crop-wild sunflower hybrids

The strength and extent of gene flow from crops into wild populations depends, in part, on the fitness of the crop alleles, as well as that of alleles at linked loci. Interest in crop-wild gene flow has increased with the advent of transgenic plants, but nontransgenic crop-wild hybrids can provide case studies to understand the factors influencing introgression, provided that the genetic architecture and the fitness effects of loci are known. This study used recombinant inbred lines (RILs) generated from a cross between crop and wild sunflowers to assess selection on domestication traits and quantitative trait loci (QTL) in two contrasting environments, in Indiana and Nebraska, USA. Only a small fraction of plants (9%) produced seed in Nebraska, due to adverse weather conditions, while the majority of plants (79%) in Indiana reproduced. Phenotypic selection analysis found that a mixture of crop and wild traits were favoured in Indiana (i.e. had significant selection gradients), including larger leaves, increased floral longevity, larger disk diameter, reduced ray flower size and smaller achene (seed) mass. Selection favouring early flowering was detected in Nebraska. QTLs for fitness were found at the end of linkage groups six (LG6) and nine (LG9) in both field sites, each explaining 11-12% of the total variation. Crop alleles were favoured on LG9, but wild alleles were favoured on LG6. QTLs for numerous domestication traits overlapped with the fitness QTLs, including flowering date, achene mass, head number, and disk diameter. It remains to be seen if these QTL clusters are the product of multiple linked genes, or individual genes with pleiotropic effects. These results indicate that crop trait values and alleles may sometimes be favoured in a noncrop environment and across broad geographical regions.     

Unconscious selection and the evolution of domesticated Plants

Two types of selection operate (and complement each other) in plants under domestication: (a) conscious or intentional selection applied by the growers for traits of interest to them; (b) unconscious or automatic selection brought about by the fact that the plants concerned were taken from their original wild habitats and placed in new (and usually very different) human-made or human-managed environments. The shift in the ecology led automatically to drastic changes in selection pressures. Numerous adaptations vital for survival in the wild environments lost their fitness under the new sets of conditions. New traits were automatically selected, resulting in the build-up of characteristic “domestication syndromes,” each fitting the specific agricultural environment provided by the farmer. The present paper assesses the evolutionary consequences of the introduction of the wild plants into several sets of contrasting farming situations. These include: (a) the type of maintenance applied, whether seed planting or vegetative propagation; (b) the plant organs for which the crop has been grown, whether they are reproductive parts or vegetative parts; (c) the impact of the system of tilling, sowing, and reaping on the evolution of grain crops; (d) the impact of the horticultural environment on fruit crops.     

Parallel domestication, convergent evolution and duplicated gene recruitment in allopolyploid cotton

A putative advantage of allopolyploidy is the possibility of differential selection of duplicated (homeologous) genes originating from two different progenitor genomes. In this note we explore this hypothesis using a high throughput, SNP-specific microarray technology applied to seed trichomes (cotton) harvested from three developmental time points in wild and modern accessions of two independently domesticated cotton species, Gossypium hirsutum and G. barbadense. We show that homeolog expression ratios are dynamic both developmentally and over the several-thousand-year period encompassed by domestication and crop improvement, and that domestication increased the modulation of homeologous gene expression. In both species, D-genome expression was preferentially enhanced under human selection pressure, but for nonoverlapping sets of genes for the two independent domestication events. Our data suggest that human selection may have operated on different components of the fiber developmental genetic program in G. hirsutum and G. barbadense, leading to convergent rather than parallel genetic alterations and resulting morphology.     

Plant Domestication and Crop Evolution in the Near East: On Events and Processes

Reconstructing the evolutionary history of crop plants is fundamental for understanding their adaptation profile and the genetic basis of yield-limiting factors, which in turn are critical for future crop improvement. A major topic in this field is the recent claim for a millennia-long ‘protracted’ domestication process. Here we evaluate the evidence for the protracted domestication model in light of published archaeobotanical data, experimental evidence and the biology of the Near Eastern crops and their wild progenitors. The crux of our discussion is the differentiation between events or ‘domestication episodes’ and the later following crop evolutionary processes under domestication (frequently termed ‘crop improvement stage’), which are by definition, still ongoing. We argue that by assuming a protracted millennia-long domestication process, one needlessly opts to operate within an intellectual framework that does not allow differentiating between the decisive (critical) domestication traits and their respective loci, and those that have evolved later during the crop dissemination and improvement following the episodic domestication event. Therefore, in our view, apart from the lack of experimental evidence to support it, the protracted domestication assumption undermines the resolution power of the study of both plant domestication and crop evolution, from the cultural as well as from the biological perspectives.     

Selection on domestication traits and quantitative trait loci in crop–wild sunflower hybrids

The strength and extent of gene flow from crops into wild populations depends, in part, on the fitness of the crop alleles, as well as that of alleles at linked loci. Interest in crop–wild gene flow has increased with the advent of transgenic plants, but nontransgenic crop–wild hybrids can provide case studies to understand the factors influencing introgression, provided that the genetic architecture and the fitness effects of loci are known. This study used recombinant inbred lines (RILs) generated from a cross between crop and wild sunflowers to assess selection on domestication traits and quantitative trait loci (QTL) in two contrasting environments, in Indiana and Nebraska, USA. Only a small fraction of plants (9%) produced seed in Nebraska, due to adverse weather conditions, while the majority of plants (79%) in Indiana reproduced. Phenotypic selection analysis found that a mixture of crop and wild traits were favoured in Indiana (i.e. had significant selection gradients), including larger leaves, increased floral longevity, larger disk diameter, reduced ray flower size and smaller achene (seed) mass. Selection favouring early flowering was detected in Nebraska. QTLs for fitness were found at the end of linkage groups six (LG6) and nine (LG9) in both field sites, each explaining 11–12% of the total variation. Crop alleles were favoured on LG9, but wild alleles were favoured on LG6. QTLs for numerous domestication traits overlapped with the fitness QTLs, including flowering date, achene mass, head number, and disk diameter. It remains to be seen if these QTL clusters are the product of multiple linked genes, or individual genes with pleiotropic effects. These results indicate that crop trait values and alleles may sometimes be favoured in a noncrop environment and across broad geographical regions.     

The role of domestication and maternal effects on seed traits of crop–wild sunflower hybrids (Helianthus annuus)

Hybridisation between crops and their wild relatives may promote the evolution of weeds. Seed germination and dormancy are the earliest life‐history traits and are highly influenced by the maternal parent. However, the ecological role of the maternal effect on seed traits in the evolution of crop–wild hybrids has received little attention. In this study, we test the relative importance of maternal and hybridisation effects on seed traits of the first generation of crop–wild sunflower hybrids (Helianthus annuus). Seed germination was tested in two wild populations with contrasting dormancy, two cultivated materials and their reciprocal crosses at four different times after harvest and three different temperatures. Seed germination at each of the four times, after ripening response and secondary dormancy were recorded along with four morphological traits. Additionally, the pericarp anatomy was analysed with light and scanning electron microscopy. We observed strong maternal effects on all seed traits. Seed germination, morphology and pericarp anatomy differed largely between the crop and wild seeds and these traits in the crop–wild hybrids resembled their female parent. Slight but significant hybridisation effects were observed in germination, mainly in seeds produced on wild plants. Crop hybridisation changed seed germination, the after ripening response and secondary dormancy in the crop direction. Morphological and anatomical traits associated with domestication strongly correlated with the observed differences in seed germination and dormancy in crop–wild sunflower hybrids. The large maternal effects along with the evolutionary divergence in seed traits were responsible for the large phenotypic differences observed in crop–wild hybrids with the same genetic composition. Wild‐like seed traits of hybrids suggest that there are no barriers to crop gene introgression at the seed level whereas crop‐like seed traits could be strongly selected against, conditioning the selection of traits expressed later in the life cycle and in the next generations.     

驯化对作物微生物组多样性和群落结构的影响及作用途径

植物与共存微生物的相互作用对植物的生长、发育、健康等具有重大影响。人类驯化导致现代作物品种与其野生祖先在生理遗传特性、生长环境等方面存在明显差异, 这必然会影响作物与其微生物组的相互作用。理解驯化对作物微生物组的影响及其作用机理, 是充分应用微生物组进行作物改良或人工育种的重要理论基础。结合课题组前期研究基础, 该文综述了驯化对作物地下和地上部分细菌和真菌(尤其是益生菌和病原菌)群落组成和多样性影响的研究现状; 并结合驯化对作物植株形态、根系构型、根系分泌物等生理特征以及生长环境的影响, 分析了驯化塑造作物微生物组的作用途径, 提出了该领域值得重点关注的研究和发展方向。     

Shifts and disruptions in resource-use trait syndromes during the evolution of herbaceous crops

Trait-based ecology predicts that evolution in high-resource agricultural environments should select for suites of traits that enable fast resource acquisition and rapid canopy closure. However, crop breeding targets specific agronomic attributes rather than broad trait syndromes. Breeding for specific traits, together with evolution in high-resource environments, might lead to reduced phenotypic integration, according to predictions from the ecological literature. We provide the first comprehensive test of these hypotheses, based on a trait-screening programme of 30 herbaceous crops and their wild progenitors. During crop evolution plants became larger, which enabled them to compete more effectively for light, but they had poorly integrated phenotypes. In a subset of six herbaceous crop species investigated in greater depth, competitiveness for light increased during early plant domestication, whereas diminished phenotypic integration occurred later during crop improvement. Mass-specific leaf and root traits relevant to resource-use strategies (e.g. specific leaf area or tissue density of fine roots) changed during crop evolution, but in diverse and contrasting directions and magnitudes, depending on the crop species. Reductions in phenotypic integration and overinvestment in traits involved in competition for light may affect the chances of upgrading modern herbaceous crops to face current climatic and food security challenges.