Domestication Traits Research Articles

Domestication During Restoration: Unintentional Selection During Eight Generations of Wild Seed Propagation Reduces Herkogamy, Dichogamy and Heterozygosity in Clarkia pulchella.

Seed production on native seed farms has increased to meet the rising demand for plant material for restoration. Although these propagation efforts are necessary for restoration, cultivating wild populations may also result in unintentional selection and elicit evolutionary changes that mimic crop domestication, essentially turning these efforts into artificial domestication experiments. Here, we investigated whether phenotypic and genomic changes associated with domestication occurred in the wildflower Clarkia pulchella Pursh (Onagraceae) by comparing the wild source populations to the farmed population after eight generations of cultivation. At the phenotypic level, the farmed population shifted towards a floral morphology associated with self-pollination, with a significant decrease in both dichogamy and herkogamy. At the genomic level, > 6500 SNPs revealed that mean expected heterozygosity of the farmed population was significantly lower than the wild populations, despite the fact that the farmed population originated from a pool of multiple wild populations. Both the shift towards a selfing phenotype and the loss of diversity are expected consequences of domestication, although the phenotypic shifts in particular occurred much more rapidly than has been observed for other domestication traits. We discuss these results in the context of plant domestication and the implications for retaining the genetic integrity of wild populations during the process of seed production for restoration.

Niger River Water Conflicts: Explanations and Forecasts

The Niger River basin home to more than 100 million people invites rigorous techniques to evaluate water conflicts among riparian areas. Conflict categories of trigger, casualty, and weapon in the interval of 2000-2022 are associated with riparian areas, revealing alternative results. The correspondence analysis reveals a relationship between riparian areas and conflicts depending on whether Benin and Niger are involved in no conflicts in the basin. The inclusion of some riparian areas in the same category implies that riparian areas with different domestic traits, such as population size, political regime, political stability, the number and length of shared borders, and the intensity of scarcity of water are similar in terms of conflicts. Future extensions of the analysis will include alternative factors, such as the destruction of nature, water pollution, and population growth, to assess water conflicts in the Niger River basin. Adding dimensions would help assess the future risks of water conflicts.

Whole-genome resequencing to investigate the genetic diversity and mechanisms of plateau adaptation in Tibetan sheep

IntroductionTibetan sheep, economically important animals on the Qinghai–Tibet Plateau, have diversified into numerous local breeds with unique characteristics through prolonged environmental adaptation and selective breeding. However, most current research focuses on one or two breeds, and lacks a comprehensive representation of the genetic diversity across multiple Tibetan sheep breeds. This study aims to fill this gap by investigating the genetic structure, diversity and high-altitude adaptation of 6 Tibetan sheep breeds using whole-genome resequencing data.ResultsSix Tibetan sheep breeds were investigated in this study, and whole-genome resequencing data were used to investigate their genetic structure and population diversity. The results showed that the 6 Tibetan sheep breeds exhibited distinct separation in the phylogenetic tree; however, the levels of differentiation among the breeds were minimal, with extensive gene flow observed. Population structure analysis broadly categorized the 6 breeds into 3 distinct ecological types: plateau-type, valley-type and Euler-type. Analysis of unique single-nucleotide polymorphisms (SNPs) and selective sweeps between Argali and Tibetan sheep revealed that Tibetan sheep domestication was associated primarily with sensory and signal transduction, nutrient absorption and metabolism, and growth and reproductive characteristics. Finally, comprehensive analysis of selective sweep and transcriptome data suggested that Tibetan sheep breeds inhabiting different altitudes on the Qinghai–Tibet Plateau adapt by enhancing cardiopulmonary function, regulating body fluid balance through renal reabsorption, and modifying nutrient digestion and absorption pathways.ConclusionIn this study, we investigated the genetic diversity and population structure of 6 Tibetan sheep breeds in Qinghai Province, China. Additionally, we analyzed the domestication traits and investigated the unique adaptation mechanisms residing varying altitudes in the plateau region of Tibetan sheep. This study provides valuable insights into the evolutionary processes of Tibetan sheep in extreme environments. These findings will also contribute to the preservation of genetic diversity and offer a foundation for Tibetan sheep diversity preservation and plateau animal environmental adaptation mechanisms.

Paralogous Gene Recruitment in Multiple Families Constitutes Genetic Architecture and Robustness of Pod Dehiscence in Legumes.

Pod dehiscence facilitates seed dispersal in wild legumes while indehiscence is a key domestication trait in cultivated ones. However, the evolutionary genetic mechanisms underlying its diversity are largely unclear. In this study, we compared transcriptomes of two warm-season (Glycine spp. and Phaseolus spp.) and two cool-season (Pisum spp. and Medicago ruthenica) legumes in analysis of dehiscent and indehiscent pod genotypes. Differentially expressed genes in AP2/ERF-like transcription factors and seven structural gene families, including lactoperoxidase, laccase, and cellulose synthase-interactive proteins, which are involved in secondary cell wall component accumulation, were identified to exert key roles in pod dehiscence variation. In accordance with this, higher lignin and cellulose contents were observed in pod secondary cell wall of dehiscent accessions of soybean and pea; however, the variation patterns of lignin polymers in soybean (accumulation) and pea (proportion) differed between dehiscent and indehiscent pods. Moreover, genome-wide comparative analysis revealed that orthogroups represented <1% of all identified differentially expressed genes could be traced among the four genera of legumes, while recruiting paralogous members may constitute the genetic robustness of legume pod dehiscence. This study compared the genetic mechanism among several legumes in pod dehiscence formation and revealed a compensating role of paralogous redundancy of involved gene families in seed dispersal, which can guide crop breeding.

The near-complete genome assembly of hexaploid wild oat reveals its genome evolution and divergence with cultivated oats.

Avena sterilis, the ancestral species of cultivated oats, is a valuable genetic resource for oat improvement. Here we generated a near-complete 10.99 Gb A. sterilis genome and a high-quality 10.89 Gb cultivated oat genome. Genome evolution analysis revealed the centromeres dynamic and structural variations landscape associated with domestication between wild and cultivated oats. Population genetic analysis of 117 wild and cultivated oat accessions worldwide detected many candidate genes associated with important agronomic traits for oat domestication and improvement. Remarkably, a large fragment duplication from chromosomes 4A to 4D harbouring many agronomically important genes was detected during oat domestication and was fixed in almost all cultivated oats from around the world. The genes in the duplication region from 4A showed significantly higher expression levels and lower methylation levels than the orthologous genes located on 4D in A. sterilis. This study provides valuable resources for evolutionary and functional genomics and genetic improvement of oat.

Telomere-to-telomere genome assembly of a male goat reveals variants associated with cashmere traits

A complete goat (Capra hircus) reference genome enhances analyses of genetic variation, thus providing insights into domestication and selection in goats and related species. Here, we assemble a telomere-to-telomere (T2T) gap-free genome (2.86 Gb) from a cashmere goat (T2T-goat1.0), including a Y chromosome of 20.96 Mb. With a base accuracy of >99.999%, T2T-goat1.0 corrects numerous genome-wide structural and base errors in previous assemblies and adds 288.5 Mb of previously unresolved regions and 446 newly assembled genes to the reference genome. We sequence the genomes of five representative goat breeds for PacBio reads, and use T2T-goat1.0 as a reference to identify a total of 63,417 structural variations (SVs) with up to 4711 (7.42%) in the previously unresolved regions. T2T-goat1.0 was applied in population analyses of global wild and domestic goats, which revealed 32,419 SVs and 25,397,794 SNPs, including 870 SVs and 545,026 SNPs in the previously unresolved regions. Also, our analyses reveal a set of selective variants and genes associated with domestication (e.g., NKG2D and ABCC4) and cashmere traits (e.g., ABCC4 and ASIP).

Identification and segregation of two closely linked major QTLs for kernel row number in advanced maize-teosinte populations.

Two closely linked novel loci, qKRN2-1 and qKRN2-2, associated with kernel row number were fine-mapped on chromosome 2, and a key candidate gene for qKRN2-1 was identified through expression analysis. Kernel row number (KRN) is a crucial factor influencing maize yield and serves as a significant target for maize breeding. The use of wild progenitor species can aid in identifying the essential traits for domestication and breeding. In this study, teosinte (MT1) served as the donor parent, the inbred maize line of Mo17 was used as the recurrent parent, we identified a major quantitative trait locus (QTL) for KRN, designated qKRN2, into two closely linked loci, qKRN2-1 and qKRN2-2. Here, fine mapping was performed to investigate two QTLs, qKRN2-1 and qKRN2-2, within a genomic range of 272kb and 775kb, respectively. This was achieved using a progeny test strategy in an advanced backcross population, with the two QTLs explaining 33.49% and 35.30% of the phenotypic variance. Molecular marker-assisted selection resulted in the development of two nearly isogenic lines (NILs), qKRN2-1 and qKRN2-2, which differed only in the segment containing the QTL. Notably, the maize (Mo17) alleles increased the KRN relative to teosinte by approximately 1.4 and 1.2 rows for qKRN2-1 and qKRN2-2, respectively. Zm00001d002989 encodes a cytokinin oxidase/dehydrogenase and its expression in the immature ears exhibited significant differences among the qKRN2-1 NILs. In situ hybridization localized Zm00001d002989 to the primordia of the inflorescence meristem and spikelet pair meristems, is predicted to be the causal gene of qKRN2-1. The findings of this study deepen our understanding of the genetic basis of KRN and hold significant potential for improving maize grain yields.

A quest for the potato of the future: Characterization of wild tuber-bearing Solanum species for de novo domestication.

The potato (Solanum tuberosum) is a staple food worldwide, but modern potato cultivation relies heavily on the use of pesticides to control pests and diseases. However, many wild Solanum species are highly resistant to biotic and abiotic stresses relevant for potato production. Several of those species have been used in potato breeding to confer resistances which has only been moderately successful. Instead, we propose an alternative approach to utilize the potential of wild Solanum germplasm. Recently, de novo domestication has been suggested to create more resilient crops: Instead of introducing resistance genes into existing crops, domestication traits could be introduced into resistant wild crop relatives to create new crops. Therefore, we selected ten promising species from the 107 known wild tuber-bearing Solanum species for their resistance to biotic and abiotic stresses based on existing scientific literature for experimental characterization looking at tuberization under short and long-day conditions, tuber glycoalkaloid content, starch digestibility and performance in tissue culture. Based on this, the highly pest and disease resistant S. bulbocastanaum was chosen. Our results showed that it produced relatively large tubers, also under long-day conditions and performed exceptionally well in tissue culture.

Pan-genome analysis reveals genomic variations during enoki mushroom domestication, with emphasis on genetic signatures of cap color and stipe length

IntroductionThe domestication of edible mushrooms, including Flammulina filiformis, offers valuable insights into the genetic changes driven by artificial selection. Understanding these changes is crucial for uncovering the mechanisms behind genome evolution in domesticated mushrooms. ObjectivesThis study aims to investigate the population structure, genetic diversity, and domestication-related genomic changes in F. filiformis. By comparing the genome sequences of 199 wild and cultivated strains, we aim to elucidate the impact of domestication on F. filiformis. MethodsWe performed de novo genome assembly and gene-based pan-genome analysis on the 199 strains, which included both wild and cultivated strains. We also conducted genome-wide association studies (GWAS) using presence-absence variation (PAV) and SNP data, combined with RNA sequencing, to identify genes associated with domestication traits, such as cap color and stipe length. Gene functional confirmation was achieved through genetic transformation experiments. ResultsOur analysis grouped the strains into four distinct populations, which correlated with varying intensities of artificial selection. The three cultivated populations exhibited smaller genome sizes, fewer genes, lower genetic variation, reduced gene expression diversity, and lower heterozygosity compared to the wild population. The analysis revealed the loss of genes related to the beta-lactam antibiotic catabolic process and specific MAPK pathway genes during domestication, rendering domesticated strains more susceptible to diseases. Four genes closely associated with cap color and stipe length were identified, but genetic transformation experiments confirmed the functional relevance of only two (FfB and FfD) identified through PAV-based GWAS. ConclusionThis study uncovered significant genomic variations between cultivated and wild F. filiformis populations, including the loss of pathogen resistance genes during domestication. We also identified key genes linked to cap color and stipe length, demonstrating for the first time the important role of PAV variation in mushroom domestication. These insights provide a foundation for future mushroom breeding and evolutionary research.

Analyzes of pan-genome and resequencing atlas unveil the genetic basis of jujube domestication

Jujube (Ziziphus jujuba Mill.), belonging to the Rhamnaceae family, is gaining increasing prominence as a perennial fruit crop with significant economic and medicinal values. Here, we conduct de novo assembly of four reference-grade genomes, encompassing one wild and three cultivated jujube accessions. We present insights into the population structure, genetic diversity, and genomic variations within a diverse collection of 1059 jujube accessions. Analyzes of the jujube pan-genome, based on our four assemblies and four previously released genomes, reveal extensive genomic variations within domestication-associated regions, potentially leading to the discovery of a candidate gene that regulates flowering and fruit ripening. By leveraging the pan-genome and a large-scale resequencing population, we identify two candidate genes involved in domestication traits, including the seed-setting rate, the bearing-shoot length and the leaf size in jujube. These genomic resources will accelerate evolutionary and functional genomics studies of jujube.

Artificial selection of mutations in two nearby genes gave rise to shattering resistance in soybean

Resistance to pod shattering is a key domestication-related trait selected for seed production in many crops. Here, we show that the transition from shattering in wild soybeans to shattering resistance in cultivated soybeans resulted from selection of mutations within the coding sequences of two nearby genes - Sh1 and Pdh1. Sh1 encodes a C2H2-like zinc finger transcription factor that promotes shattering by repressing SHAT1-5 expression, thereby reducing the secondary wall thickness of fiber cap cells in the abscission layers of pod sutures, while Pdh1 encodes a dirigent protein that orchestrates asymmetric lignin distribution in inner sclerenchyma, creating torsion in pod walls that facilitates shattering. Integration analyses of quantitative trait locus mapping, genome-wide association studies, and allele distribution in representative soybean germplasm suggest that these two genes are primary modulators underlying this domestication trait. Our study thus provides comprehensive understanding regarding the genetic, molecular, and cellular bases of shattering resistance in soybeans.

Improving complex agronomic and domestication traits in the perennial grain crop intermediate wheatgrass with genetic mapping and genomic prediction.

The perennial grass Thinopyrum intermedium (intermediate wheatgrass [IWG]) is being domesticated as a food crop. With a deep root system and high biomass, IWG can help reduce soil and water erosion and limit nutrient runoff. As a novel grain crop undergoing domestication, IWG lags in yield, seed size, and other agronomic traits compared to annual grains. Better characterization of trait variation and identification of genetic markers associated with loci controlling the traits could help in further improving this crop. The University of Minnesota's Cycle 5 IWG breeding population of 595 spaced plants was evaluated at two locations in 2021 and 2022 for agronomic traits plant height, grain yield, and spike weight, and domestication traits shatter resistance, free grain threshing, and seed size. Pairwise trait correlations were weak to moderate with the highest correlation observed between seed size and height (0.41). Broad-sense trait heritabilities were high (0.68-0.77) except for spike weight (0.49) and yield (0.44). Association mapping using 24,284 genome-wide single nucleotide polymorphism markers identified 30 main quantitative trait loci (QTLs) across all environments and 32 QTL-by-environment interactions (QTE) at each environment. The genomic prediction model significantly improved predictions when parents were used in the training set and significant QTLs and QTEs used as covariates. Seed size was the best predicted trait with model predictive ability (r) of 0.72; yield was predicted moderately well (r=0.45). We expect this discovery of significant genomic loci and mostly high trait predictions from genomic prediction models to help improve future IWG breeding populations.

Irregular green netting of eggplant fruit peel: a domestication trait controlled by SmGLK2 with potential for fruit colour diversification.

The distribution of chlorophylls in eggplant (Solanum melongena) peel exhibits either a uniform pattern or an irregular green netting pattern. The latter, manifested as a gradient of dark green netting that is intensified in the proximal part of the fruit on a pale green background, is common in wild relatives and some eggplant landraces. Despite the selection of uniform chlorophylls during domestication, the netting pattern contributes to a greater diversity of fruit colours. Here, we used over 2300 individuals from different populations, including a multi-parent advanced generation inter-cross population for candidate genomic region identification, an F2 population for bulked segregant analysis by sequencing, and advanced backcrosses for edges-to-core fine-mapping, to identify SmGLK2 gene as responsible for the irregular netting in eggplant fruits. We also analysed the gene sequence of 178 S. melongena accessions and 22 wild relative species for tracing the evolutionary changes that the gene has undergone during domestication. Three different mutations were identified leading to the absence of netting. The main causative indel induces a premature stop codon disrupting the protein conformation and function, which was confirmed by western blot analysis and confocal microscopy observations. SmGLK2 has a major role in regulating chlorophyll biosynthesis in eggplant fruit peel.

Pan-genome analysis of 13 Spinacia accessions reveals structural variations associated with sex chromosome evolution and domestication traits in spinach.

Structural variations (SVs) are major genetic variants that can be involved in the origin, adaptation and domestication of species. However, the identification and characterization of SVs in Spinacia species are rare due to the lack of a pan-genome. Here, we report eight chromosome-scale assemblies of cultivated spinach and its two wild species. After integration with five existing assemblies, we constructed a comprehensive Spinacia pan-genome and identified 193 661 pan-SVs, which were genotyped in 452 Spinacia accessions. Our pan-SVs enabled genome-wide association study identified signals associated with sex and clarified the evolutionary direction of spinach. Most sex-linked SVs (86%) were biased to occur on the Y chromosome during the evolution of the sex-linked region, resulting in reduced Y-linked gene expression. The frequency of pan-SVs among Spinacia accessions further illustrated the contribution of these SVs to domestication, such as bolting time and seed dormancy. Furthermore, compared with SNPs, pan-SVs act as efficient variants in genomic selection (GS) because of their ability to capture missing heritability information and higher prediction accuracy. Overall, this study provides a valuable resource for spinach genomics and highlights the potential utility of pan-SV in crop improvement and breeding programmes.

Selection against domestication alleles in introduced rabbit populations.

Humans have moved domestic animals around the globe for thousands of years. These have occasionally established feral populations in nature, often with devastating ecological consequences. To understand how natural selection shapes re-adaptation into the wild, we investigated one of the most successful colonizers in history, the European rabbit. By sequencing the genomes of 297 rabbits across three continents, we show that introduced populations exhibit a mixed wild-domestic ancestry. We show that alleles that increased in frequency during domestication were preferentially selected against in novel natural environments. Interestingly, causative mutations for common domestication traits sometimes segregate at considerable frequencies if associated with less drastic phenotypes (for example, coat colour dilution), whereas mutations that are probably strongly maladaptive in nature are absent. Whereas natural selection largely targeted different genomic regions in each introduced population, some of the strongest signals of parallelism overlap genes associated with neuronal or brain function. This limited parallelism is probably explained by extensive standing genetic variation resulting from domestication together with the complex mixed ancestry of introduced populations. Our findings shed light on the selective and molecular mechanisms that enable domestic animals to re-adapt to the wild and provide important insights for the mitigation and management of invasive populations.

Domestication as the evolution of interspecies cooperative breeding.

We propose that domestication is the result of interspecies cooperative breeding. Considering domestication as an outcome of cooperative breeding can explain how domestication occurs in both plants and animals, encompass cases of domestication that do not involve humans, and shed light on why humans are involved in so many domesticatory relationships. We review the cooperative breeding model of human evolution, which posits that care of human infants by alloparents enabled the evolution of costly human brains and long juvenile development, while selecting for tolerance of strangers. We then explore how human cooperation in the protection and provisioning of young plants and animals can explain the evolution of domestication traits such as changes in development; loss of aggressive, defensive, and bet-hedging aspects of the phenotype; and increased fertility. We argue that the importance of cooperative breeding to human societies has made humans especially likely to enter into interspecies cooperative breeding relationships.

Engineering the future of Physalis grisea: A focus on agricultural challenges, model species status, and applied improvements

Societal Impact StatementGroundcherry (Physalis grisea) is a plant species grown for its flavorful fruit. The fruit drops from the plant, hence the common name groundcherry. This makes harvest cumbersome and puts the fruit at risk for carrying soil‐borne pathogens, therefore making them unsellable. Furthermore, insects often damage the plants, reducing yield. Advances in gene editing offer promise for addressing these issues and aiding home gardeners and farmers. Improvement will expand access to this nutritious fruit, rich in potassium, vitamin C, and antioxidants. Additionally, studies of its biology could serve as a model for improving other fruiting plants, particularly underutilized species.SummaryP. grisea is an underutilized, semidomesticated fruit crop with rising agronomic value. Several resources have been developed for its use in fundamental biological research, including a plant transformation system and a high‐quality reference genome. Already, P. grisea has been used as a model to investigate biological phenomena including inflated calyx syndrome and gene compensation. P. grisea has also been used to demonstrate the potential of fast‐tracking domestication trait improvement through approaches such as CRISPR/Cas9 gene editing. This work has led to the Physalis Improvement Project, which relies on reverse genetics to understand the mechanisms that underlie fruit abscission and plant–herbivore interactions to guide approaches for improvement of undesirable characteristics. CRISPR/Cas9 gene editing has been used to target P. grisea genes that are suspected to act in fruit abscission, particularly orthologs of those that are reported in tomato abscission zone development. A similar approach is being taken to target P. grisea genes involved in the withanolide biosynthetic pathway to determine the impact of withanolides on plant–herbivore interactions. Results from these research projects will lead to a greater understanding of important biological processes and will also generate knowledge needed to develop cultivars with reduced fruit drop and increased resistance to insect herbivory.

Analysis of goat PPP6C mRNA profile, detection of genetic variations, and their associations with litter size

The Protein Phosphatase 6 Catalytic Subunit (PPP6C) is evolutionarily a conserved gene in eukaryotes known to play a significant role in mammalian reproduction. This study aimed to investigate expression patterns of PPP6C and explore its association with litter size in Shaanbei white cashmere (SBWC) goats. Initially, we determined the mRNA expression levels of PPP6C in both male and female goats across multiple tissues. The results showed that PPP6C mRNA was expressed in multiple tissues, with higher levels in the testis and fallopian tubes, suggesting its involvement in goat reproduction. Additionally, we identified a novel 19 bp InDel within the PPP6C gene in a population of 1030 SBWC goats, which exhibited polymorphism. Statistical analysis revealed a significant association between the19 bp InDel mutation and litter size (P < 0.05). Subsequent, bioinformatics analysis, including linkage disequilibrium (LD) block and selective scanning, highlighted the linkage tendency among most InDel loci did not stand out within B-8 block, there were still some InDel loci linked to the 19 bp within a relatively narrow region. Furthermore, comparative analysis with Bezoars, these selective signals all indicated that this gene was under higher selection pressure, implying that the 19 bp InDel locus within the PPP6C is potentially associated with domesticated traits, particularly in relation to litter size. The results of the present study suggest that the PPP6C is a vital candidate gene affecting prolificacy in goats, with implications for selective breeding programs for goat breeds.

A pan-TE map highlights transposable elements underlying domestication and agronomic traits in Asian rice.

Transposable elements (TEs) are ubiquitous genomic components and hard to study due to being highly repetitive. Here we assembled 232 chromosome-level genomes based on long-read sequencing data. Coupling the 232 genomes with 15 existing assemblies, we developed a pan-TE map comprising both cultivated and wild Asian rice. We detected 177084 high-quality TE variations and inferred their derived state using outgroups. We found TEs were one source of phenotypic variation during rice domestication and differentiation. We identified 1246 genes whose expression variation was associated with TEs but not single-nucleotide polymorphisms (SNPs), such as OsRbohB, and validated OsRbohB's relative expression activity using a dual-Luciferase (LUC) reporter assays system. Our pan-TE map allowed us to detect multiple novel loci associated with agronomic traits. Collectively, our findings highlight the contributions of TEs to domestication, differentiation and agronomic traits in rice, and there is massive potential for gene cloning and molecular breeding by the high-quality Asian pan-TE map we generated.

Long noncoding RNAs underlie multiple domestication traits and leafhopper resistance in soybean.

The origin and functionality of long noncoding RNA (lncRNA) remain poorly understood. Here, we show that multiple quantitative trait loci modulating distinct domestication traits in soybeans are pleiotropic effects of a locus composed of two tandem lncRNA genes. These lncRNA genes, each containing two inverted repeats, originating from coding sequences of the MYB genes, function in wild soybeans by generating clusters of small RNA (sRNA) species that inhibit the expression of their MYB gene relatives through post-transcriptional regulation. By contrast, the expression of lncRNA genes in cultivated soybeans is severely repressed, and, consequently, the corresponding MYB genes are highly expressed, shaping multiple distinct domestication traits as well as leafhopper resistance. The inverted repeats were formed before the divergence of the Glycine genus from the Phaseolus-Vigna lineage and exhibit strong structure-function constraints. This study exemplifies a type of target for selection during plant domestication and identifies mechanisms of lncRNA formation and action.