More than one way to evolve a weed: parallel evolution of US weedy rice through independent genetic mechanisms

Many different crop species were selected for a common suite of ‘domestication traits’, which facilitates their use for studies of parallel evolution. Within domesticated rice (Oryza sativa), there has also been independent evolution of weedy strains from different cultivated varieties. This makes it possible to examine the genetic basis of parallel weed evolution and the extent to which this process occurs through shared genetic mechanisms. We performed comparative QTL mapping of weediness traits using two recombinant inbred line populations derived from crosses between an indica crop variety and representatives of each of the two independently evolved weed strains found in US rice fields, strawhull (S) and blackhull awned (B). Genotyping‐by‐sequencing provided dense marker coverage for linkage map construction (average marker interval <0.25 cM), with 6016 and 13 730 SNPs mapped in F₅ lines of the S and B populations, respectively. For some weediness traits (awn length, hull pigmentation and pericarp pigmentation), QTL mapping and sequencing of underlying candidate genes confirmed that trait variation was largely attributable to individual loci. However, for more complex quantitative traits (including heading date, panicle length and seed shattering), we found multiple QTL, with little evidence of shared genetic bases between the S and B populations or across previous studies of weedy rice. Candidate gene sequencing revealed causal genetic bases for 8 of 27 total mapped QTL. Together these findings suggest that despite the genetic bottleneck that occurred during rice domestication, there is ample genetic variation in this crop to allow agricultural weed evolution through multiple genetic mechanisms.     

REDUCED DROUGHT TOLERANCE DURING DOMESTICATION AND THE EVOLUTION OF WEEDINESS RESULTS FROM TOLERANCE–GROWTH TRADE‐OFFS

The increased reproductive potential, size, shoot allocation, and growth rate of weedy plants may result from reduced resource allocation to other aspects of plant growth and defense. To investigate whether changes in resource allocation occurred during domestication or the evolution of weediness, we compared the mycorrhizal responsiveness, growth, and drought tolerance of nine native ruderal, nine agriculturally weedy (four U.S. weedy and five Australian weedy), and 14 domesticated populations (eight ancient landraces and six improved cultivars) of the common sunflower (Helianthus annuus). Domesticated sunflower cultivars were less drought tolerant, but had higher plant growth and fecundity and coarser roots than wild populations. There were no changes in level of drought tolerance between improved cultivars and ancient landrace plants, but there was an increase in allocation to flowers with recent selection. Weedy populations were intermediate between domesticated cultivars and native ruderal populations for plant growth rate, root architecture, and drought tolerance. Weedy populations benefited most from mycorrhizal inoculation by having fewer wilted leaves and wetter soil. Overall, we found that trade‐offs between drought tolerance and several aspects of plant growth, including growth rate, allocation to flowering, and root architecture, govern evolution during sunflower domestication and the invasion of disturbed habitat.     

Soil burial induced dormancy in weedy rice seeds through hormone level changes: Implications in adaptive evolution and weed control

Seed dormancy plays a key role in preventing seeds of higher plants from random germination under adverse environmental conditions. Previous studies suggested that a critical temperature could regulate germination of weedy rice seeds without primary dormancy at seed dispersion. However, what will happen to the non-dormant seeds after shattering in the soil seed banks when temperature fluctuates to exceed the critical temperature remains an interesting question to be investigated. To determine whether or not soil burial can change the status of dormancy in weedy rice seeds, we examined germination ratios of weedy rice seeds after soil-burial treatments. In addition, we compared hormone levels in the untreated seeds and viable but ungerminated seeds after soil burial. Results showed that soil burial induced a proportion of 41%–72% dormant seeds in the initially non-dormant weedy rice seeds. Also, the induction of seed dormancy is associated with the change of hormone levels in the seeds treated by soil burial, suggesting that soil burial can significantly activate the control of hormone production in seeds. Together, the previously reported mechanism of critical temperature-inhibited seed germination and the newly found phenomenon of soil burial-induced seed dormancy provide a “double-security” strategy to ensure germination of weedy rice seeds under a favorable condition in agricultural ecosystems. The findings not only reveal the important role of rapid evolution of adaptive functions in weeds, such as weedy rice, in adapting to changing agricultural environments, but also facilitate the design of strategies for effective weedy rice control practices.     

Genetics and evolution of weedy Helianthus annuus populations: adaptation of an agricultural weed

Agricultural weeds are a major cost to economies throughout the world, and have evolved from numerous plant species in many different plant families. Despite their ubiquity, we do not yet know how easily or often weeds evolve from their wild ancestors or the kinds of genes underlying their evolution. Here we report on the evolution of weedy populations of the common sunflower Helianthus annuus. We analysed 106 microsatellites in 48 individuals from each of six wild and four weed populations of the species. The statistical tests lnRV and lnRH were used to test for significant reductions in genetic variability at each locus in weedy populations compared to nearby wild populations. Between 1% and 6% of genes were significant outliers with reduced variation in weedy populations, implying that a small but not insignificant fraction of the genome may be under selection and involved in adaptation of weedy sunflowers. However, there did not appear to be a substantial reduction in variation across the genome, suggesting that effective population sizes have remained very large during the recent evolution of these weedy populations. Additional analyses showed that weedy populations are more closely related to nearby wild populations than to each other, implying that weediness likely evolved multiple times within the species, although a single origin followed by gene flow with local populations cannot be ruled out. Together, our results point to the relative ease with which weedy forms of this species can evolve and persist despite the potentially high levels of geneflow with nearby wild populations.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Dormancy genes from weedy rice respond divergently to seed development environments

Genes interacting with seed developmental environments control primary dormancy. To understand how a multigenic system evolved to adapt to the changing environments in weedy rice, we evaluated genetic components of three dormancy QTL in a synchronized nondormant genetic background. Two genetically identical populations segregating for qSD1, qSD7-1, and qSD12 were grown under greenhouse and natural conditions differing in temperature, relative humidity, and light intensity during seed development. Low temperatures tended to enhance dormancy in both conditions. However, genotypes responded to the environments divergently so that two populations displayed similar distributions for germination. Additive and/or dominance effects of the three loci explained approximately 90% of genetic variances and their epistases accounted for the remainder in each environment. The qSD1 and qSD7-1 main effects were increased, while the qSD12 additive effect was decreased by relatively low temperatures. Both gene main and epistatic effects were involved in G x E interactions, which in magnitude were greater than environmental main effect. The divergent responses of dormancy genes observed in this simple multigenic system presumably have selective advantages in natural populations adapted to changing environments and hence represent a genetic mechanism stabilizing the dormancy level of weedy rice ripened in different seasons or temperature regimes.     

Better performance of germination in hyperosmotic solutions in conspecific weedy rice than cultivated rice

Weeds and crops that grow together often confront similar types of environmental stress, especially drought stress. Weedy rice (Oryza sativa f. spontanea) and cultivated rice (O. sativa L.) provide a unique pair consisting of a weed and a conspecific model crop that can be used to study the drought tolerance of plants across a large distributional range. The investigation on weedy rice's damage to paddy fields showed that it was more serious in dry direct seeding than water direct seeding. Compared with water direct seeding, the seeds of cultivated rice and weedy rice in dry direct seeding will absorb water and germinate under the condition of insufficient soil moisture. Our hypothesis is that weedy rice seeds have evolved stronger germination ability than coexisting cultivated rice under water stress, so that they can obtain more growth space in the early stage in dry direct seeding and thus obtain higher fitness. Seeds of weedy rice populations and coexisting rice cultivars were collected from 61 sites across China and were germinated with 20% polyethylene glycol‐6000 to simulate drought stress. Two drought response indices, which assessed germination rate and germination index, plus one germination stress tolerance index, indicated significantly greater drought tolerance in weedy rice populations than in coexisting rice cultivars (P < 0.01). Drought tolerance for the three indexes were indica weedy rice > indica rice cultivars, japonica weedy rice > japonica rice cultivars, and indica weedy rice > japonica rice cultivars. These results indicate that weedy rice populations show stronger drought stress tolerance than coexisting rice cultivars at various sites, specifically during the seed germination period. Furthermore, Pearson's correlation found that drought response of weedy rice populations and coexisting rice cultivars were significantly different with these environmental factors: latitude, altitude, annual mean precipitation, mean annual temperature, mean precipitation in the sowing month, mean temperature in the sowing month, and sowing methods. Weedy rice shows different patterns of drought tolerance variation across geographical (latitude and altitude) and environmental (precipitation) gradients compared to coexisting rice cultivars. This study suggests that weedy rice might have evolved new drought tolerance and could provide a useful source of genetic resources for improving drought tolerance of crop cultivars and breeding direct seeded cultivars to reduce the usage of seeds in direct seeding.     

Climate change and plant regeneration from seed

At the core of plant regeneration, temperature and water supply are critical drivers for seed dormancy (initiation, break) and germination. Hence, global climate change is altering these environmental cues and will preclude, delay, or enhance regeneration from seeds, as already documented in some cases. Along with compromised seedling emergence and vigour, shifts in germination phenology will influence population dynamics, and thus, species composition and diversity of communities. Altered seed maturation (including consequences for dispersal) and seed mass will have ramifications on life history traits of plants. Predicted changes in temperature and precipitation, and thus in soil moisture, will affect many components of seed persistence in soil, e.g. seed longevity, dormancy release and germination, and soil pathogen activity. More/less equitable climate will alter geographic distribution for species, but restricted migratory capacity in some will greatly limit their response. Seed traits for weedy species could evolve relatively quickly to keep pace with climate change enhancing their negative environmental and economic impact. Thus, increased research in understudied ecosystems, on key issues related to seed ecology, and on evolution of seed traits in nonweedy species is needed to more fully comprehend and plan for plant responses to global warming.     

GERMINATION AND SEEDLING EMERGENCE IN SPERGULA ARVENSIS

Spergula arvensis L. (Caryophyllaceae) is a cosmopolitan weedy annual that exhibits a genetically based seed coat polymorphism in which two primary seed types, papillate (P) and nonpapillate (NP), are produced. This study examined germination response of the two morphs and seedling emergence in Pacific coast populations. Germination behavior was complex and variable among populations and between seed morphs. Seeds from some populations exhibited greater temperature sensitivity and/or differentiation between morphs. Freshly produced seeds from grassland and ruderal populations were strongly dormant at the time of dispersal, in contrast to seeds from cultivated or summer-wet populations. In San Francisco Bay area populations, germination and seedling emergence was largely simultaneous, following the first substantial fall rains.     

In situ management and domestication of plants in Mesoamerica

BACKGROUND AND AIMS: Ethnobotanical studies in Mexico have documented that Mesoamerican peoples practise systems of in situ management of wild and weedy vegetation directed to control availability of useful plants. In situ management includes let standing, encouraging growing and protection of individual plants of useful species during clearance of vegetation, which in some cases may involve artificial selection. The aim of this study was to review, complement and re-analyse information from three case studies which examined patterns of morphological, physiological and genetic effects of artificial selection in plant populations under in situ management in the region. METHODS: Information on wild and in situ managed populations of the herbaceous weedy plants Anoda cristata and Crotalaria pumila, the tree Leucaena esculenta subsp. esculenta and the columnar cacti Escontria chiotilla, Polaskia chichipe and Stenocereus stellatus from Central Mexico was re-analysed. Analyses compared morphology and frequency of morphological variants, germination patterns, and population genetics parameters between wild and managed in situ populations of the species studied. Species of columnar cacti are under different management intensities and their populations, including cultivated stands of P. chichipe and S. stellatus, were also compared between species. KEY RESULTS: Significant differences in morphology, germination patterns and genetic variation documented between wild, in situ managed and cultivated populations of the species studied are associated with higher frequencies of phenotypes favoured by humans in managed populations. Genetic diversity in managed populations of E. chiotilla and P. chichipe is slightly lower than in wild populations but in managed populations of S. stellatus variation was higher than in the wild. However, genetic distance between populations was generally small and influenced more by geographic distance than by management. CONCLUSIONS: Artificial selection operating on in situ managed populations of the species analysed is causing incipient domestication. This process could be acting on any of the 600-700 plant species documented to be under in situ management in Mesoamerica. In situ domestication of plants could be relevant to understand early processes of domestication and current conditions of in situ conservation of plant genetic resources.     

蔊菜杂草性的研究

本文通过对十字花科常见杂草芹菜[Rorippa in lica(L.)Hiern]在南京地区的自然群体的周期性观察,以了解其在不同生态环境下的生活周期。本文还通过一系列实验来观察其繁育系统、种子传播的动因、结实力、种子发芽率和植物抗人为干扰的耐受性。花蕾套袋试验证明蔊菜主要是自交可亲和的,即使存在异型杂交也是微不足道的。种子传播效应试验揭示水和风都是种子传播的自然力。该种具有范围很广的耐受性,踏践和刈割试验证明在严重的人为于扰下仍能完成其生活周期。蔊菜尽管本质上属于多年生草本,能产生大量的种子,在很大程度上靠种子繁殖。种子萌芽试验证明它的种子萌发参差不齐。总之,蔊菜具有典型杂草的许多特性,而这些特性给它以适合在多种自然的和人为干扰的环境中正常地生长的能力。     

Malaysian weedy rice shows its true stripes: wild Oryza and elite rice cultivars shape agricultural weed evolution in Southeast Asia

Weedy rice is a close relative of domesticated rice (Oryza sativa) that competes aggressively with the crop and limits rice productivity worldwide. Most genetic studies of weedy rice have focused on populations in regions where no reproductively compatible wild Oryza species occur (North America, Europe and northern Asia). Here, we examined the population genetics of weedy rice in Malaysia, where wild rice (O. rufipogon) can be found growing in close proximity to cultivated and weedy rice. Using 375 accessions and a combined analysis of 24 neutral SSR loci and two rice domestication genes (sh4, controlling seed shattering, and Bh4, controlling hull colour), we addressed the following questions: (i) What is the relationship of Malaysian weedy rice to domesticated and wild rice, and to weedy rice strains in the USA? (ii) To what extent does the presence of O. rufipogon influence the genetic and phenotypic diversity of Malaysian weeds? (iii) What do the distributions of sh4 and Bh4 alleles and associated phenotypes reveal about the origin and contemporary evolution of Malaysian weedy rice? Our results reveal the following: independent evolutionary origins for Malaysian weeds and US strains, despite their very close phenotypic resemblance; wild‐to‐weed gene flow in Malaysian weed populations, including apparent adaptive introgression of seed‐shattering alleles; and a prominent role for modern Malaysian cultivars in the origin and recent proliferation of Malaysian weeds. These findings suggest that the genetic complexity and adaptability of weedy crop relatives can be profoundly influenced by proximity to reproductively compatible wild and domesticated populations.     

Are plant populations seed‐limited? A review of seed sowing experiments

We define seed limitation to be an increase in population size following seed addition. Here, we briefly consider how theoretical models deal with seed limitation and how seed sowing experiments can be used to unravel the extent of seed limitation in natural systems. We review two types of seed addition experiments: seed augmentation studies where seeds are added to existing populations; and seed introductions where seeds are sown in unoccupied sites. Overall, approximately 50% of seed augmentation experiments show evidence of seed limitation. These studies show that seed limitation tends to occur more commonly in early successional habitats and in early successional species. Most of the studies have concentrated on simply categorising populations as seed- or microsite-limited, but we believe that seed sowing experiments could be used to reveal much more about community structure, and we discuss possible future directions. In 53% of introduction studies (where seeds were sown at sites from which the species was known to be absent) the introduced species was recorded in at least one of the experimental sites following sowing. However, of the subset of studies where both seedlings and adult plants were recorded, 64% of sites contained seedlings while only 23% contained adults. This implies that, for many species, conditions for establishment are more stringent than conditions for germination. The successful establishment of plants in unoccupied patches indicates the potential for immigration to enhance local diversity (the spatial mass effect). Few studies continued monitoring for long enough to determine whether or not self-sustaining populations were successfully established, and no study attempted to link introduction sites to a putative natural source of propagules, or considered the dynamics of the metapopulation as a whole.     

Molecular approaches to origin,ancestry and domestication history of crop plants: Barley and clover as examples

Knowledge of the origin and domestication history of crop plants is important for studies aiming at avoiding the erosion of genetic resources due to the loss of ecotypes and landraces and habitats and increased urbanization. Such knowledge also strengthens the capacity of modern farming system to develop and scale-up the domestication of high value potential crops that can be achieved by improving the knowledge that help to identify and select high value plant species within their locality, identify and apply the most appropriate propagation techniques for improving crops and integrate improved crop species into the farming systems. The study of domestication history and ancestry provide means for germplasm preservation through establishment of gene banks, largely as seed collections, and preservation of natural habitats. Information about crop evolution and specifically on patterns of genetic change generated by evolution prior, during, and after domestication, is important to develop sound genetic conservation programs of genetic resources of crop plants and also increases the efficiency of breeding programs. In recent years, molecular approaches have contributed to our understanding of the aspects of plant evolution and crops domestication. In this article, aspects of crops domestication are outlined and the role of molecular data in elucidating the ancestry and domestication of crop plants are outlined. Particular emphasis is given to the contribution of molecular approaches to the origin and domestication history of barley and the origin and ancestry of the Egyptian clover.     

Not all weeds are created equal: A database approach uncovers differences in the sexual system of native and introduced weeds

Weedy species provide excellent opportunities to examine the process of successful colonization of novel environments. Despite the influence of the sexual system on a variety of processes from reproduction to genetic structure, how the sexual system of species influences weediness has received only limited consideration. We examined the hypothesis that weedy plants have an increased likelihood of being self‐compatible compared with nonweedy plants; this hypothesis is derived from Baker's law, which states that species that can reproduce uniparentally are more likely to successfully establish in a new habitat where mates are lacking. We combined a database of the weed (weedy/nonweedy) and introduction status (introduced/native) of plant species found in the USA with a database of plant sexual systems and determined whether native and introduced weeds varied in their sexual systems compared with native and introduced nonweeds. We found that introduced weeds are overrepresented by species with both male and female functions present within a single flower (hermaphrodites) whereas weeds native to the USA are overrepresented by species with male and female flowers present on a single plant (monoecious species). Overall, our results show that Baker's law is supported at the level of the sexual system, thus providing further evidence that uniparental reproduction is an important component of being either a native or introduced weed.     

Genetic diversity and gene flow among pearl millet crop/weed complex: a case study

Weedy plants with intermediate (domesticated × wild) phenotypes occur in most pearl millet fields in West Africa, even in the absence of wild populations. They are usually found, in high numbers, both inside and outside of drills. Questions pertaining to the evolutionary dynamics of diversity within the pearl millet complex (domesticated–weedy–wild forms) were addressed in this study. The diversity of the different components of this complex sampled in two pearl millet fields in two villages of southwestern Niger was assessed at both molecular (AFLP) and morphological levels. Results show that, in both fields, weedy plants found outside of drills are morphologically distinct from weedy plants found inside drills, despite their close similarity at AFLP markers. The data suggest some introgression from the wild to the weedy population but nevertheless that the gene flow between the parapatric wild and domesticated populations is very low. This challenges the traditional view that regular hybridization between domesticated and wild pearl millets explains the abundance of these weedy plants despite farmers’ seed selection. The level of genetic differentiation between fields from the two villages was low when considering domesticated and weedy plants. This could be explained by high gene flow resulting from substantial seed exchanges between farmers. The fact that it is very difficult for farmers to keep their own selected seeds, and the consequent substantial seed exchanges between them, is probably the main factor accounting for the maintenance and dispersal of weedy pearl millets in the region, even in areas where no wild forms have been observed.C. Mariac and T. Robert contributed equally to the work.     

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.     

The evolutionary potential of Baker's weediness traits in the common morning glory, Ipomoea purpurea (Convolvulaceae)

? Premise of the study: Many reports have cited Baker's list of weediness traits, or those that exemplify the "ideal" weed, yet few have considered the evolutionary potential of such traits as a group. Thus, it is unknown whether constraints on the evolution of increased weediness, such as a lack of genetic variation or genetic correlations between the traits, are present. Ipomoea purpurea, the common morning glory, is a problematic weed that exhibits many of Baker's ideal weed traits. ? Methods: We used progeny from a half/full-sib breeding design in a series of three greenhouse experiments to assess the presence of genetic variation, narrow sense heritabilities, and genetic correlations in Baker's growth, competition, and fitness "weediness" traits in two populations of I. purpurea. ? Key results: We uncovered genetic variation underlying reproductive fitness traits and competitive ability in at least one population, but no evidence of genetic variation underlying growth rate in either population. Genetic correlations between many of the weediness characters differed significantly from zero; however, their direction and/or magnitude differed between populations. ? Conclusions: We found that increased weediness in the common morning glory is more likely to occur through selection on reproductive output and competitive ability rather than through selection on growth rate. Assessing Baker's traits in a quantitative genetics framework can provide a solid perspective on their evolutionary potential and a unique framework within which to determine how weeds will respond to different environmental stresses and/or scenarios of global climate change.     

All roads lead to weediness: Patterns of genomic divergence reveal extensive recurrent weedy rice origins from South Asian Oryza

Weedy rice (Oryza spp.), a weedy relative of cultivated rice (O. sativa), infests and persists in cultivated rice fields worldwide. Many weedy rice populations have evolved similar adaptive traits, considered part of the ‘agricultural weed syndrome’, making this an ideal model to study the genetic basis of parallel evolution. Understanding parallel evolution hinges on accurate knowledge of the genetic background and origins of existing weedy rice groups. Using population structure analyses of South Asian and US weedy rice, we show that weeds in South Asia have highly heterogeneous genetic backgrounds, with ancestry contributions both from cultivated varieties (aus and indica) and wild rice. Moreover, the two main groups of weedy rice in the USA, which are also related to aus and indica cultivars, constitute a separate origin from that of Asian weeds. Weedy rice populations in South Asia largely converge on presence of red pericarps and awns and on ease of shattering. Genomewide divergence scans between weed groups from the USA and South Asia, and their crop relatives are enriched for loci involved in metabolic processes. Some candidate genes related to iconic weedy traits and competitiveness are highly divergent between some weed‐crop pairs, but are not shared among all weed‐crop comparisons. Our results show that weedy rice is an extreme example of recurrent evolution, and suggest that most populations are evolving their weedy traits through different genetic mechanisms.     

Sequence polymorphisms in wild,weedy, and cultivated rice suggest seed‐shattering locus sh4 played a minor role in Asian rice domestication

The predominant view regarding Asian rice domestication is that the initial origin of nonshattering involved a single gene of large effect, specifically, the sh4 locus via the evolutionary replacement of a dominant allele for shattering with a recessive allele for reduced shattering. Data have accumulated to challenge this hypothesis. Specifically, a few studies have reported occasional seed‐shattering plants from populations of the wild progenitor of cultivated rice (Oryza rufipogon complex) being homozygous for the putative “nonshattering” sh4 alleles. We tested the sh4 hypothesis for the domestication of cultivated rice by obtaining genotypes and phenotypes for a diverse set of samples of wild, weedy, and cultivated rice accessions. The cultivars were fixed for the putative “nonshattering” allele and nonshattering phenotype, but wild rice accessions are highly polymorphic for the putative “nonshattering” allele (frequency ~26%) with shattering phenotype. All weedy rice accessions are the “nonshattering” genotype at the sh4 locus but with shattering phenotype. These data challenge the widely accepted hypothesis that a single nucleotide mutation (“G”/“T”) of the sh4 locus is the major driving force for rice domestication. Instead, we hypothesize that unidentified shattering loci are responsible for the initial domestication of cultivated rice through reduced seed shattering.