Do Hawthorn Berries Stain

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Hawthorn is of high economic value due to its medicinal properties and health benefits. Crataegus is a member of the Rosaceae family; The genus has a complex taxonomic history, and several theories have been proposed about its origin. In this study, 53 accessions from seven Crataegus taxa indigenous to China and accessions from exotic Crataegus species (two from Europe and one from North America) were analyzed by amplified in situ fragment sequencing (SLAF-seq). In total, 933, 450 single nucleotide polymorphisms were identified after filtering and used to investigate the genetic evolution of the species. Phylogenetic trees derived from Simple Nuclear Sequence Repeats (SSRs) and SLAF-seq data showed the same structure, with Crataegus maximowiczii and Crataegus sanguineae forming a closely related cluster clearly separated from the cluster consisting of Crataegus hupehensis, Crataegus pinnatifida and Crataegus pinnatifida var. major, Crataegus bretschneideri and Crataegus scabrifolia. Phylogenetic analysis indicated that the seven Chinese Crataegus taxa had two separate events of speciation. Plants that developed the southwestern pathway shared a gene pool with European species, while plants along the northeastern pathway shared a gene pool with North American species. TreeMix genetic analysis revealed that C. bretschneideri may have a hybrid origin. This study provides valuable information on the origins of the Chinese Crataegus and suggests an evolutionary model for the major Crataegus species native to China.

Do Hawthorn Berries Stain

The genus Crataegus (Hawthorn), a member of the Rosaceae family, ranges from small shrubs to trees distributed in Eurasia and America (Phipps et al., 1990). Hawthorn is among the most economically important plant species in China, due to its pleasant flavor, attractive color and nutrient-rich fruit (Xu et al., 2016). The use of hawthorn in preventive medicine dates back to the late 19th century. Hawthorn contains bioactive compounds, such as flavonoids, phenols, and oligomeric procyanidins, which have therapeutic benefits (Dickinson et al., 2014; Dahmer and Scott, 2018). Laboratory tests and previous clinical trials have demonstrated the efficacy of hawthorn in the treatment and prevention of cardiovascular disease (Edwards et al., 2012).

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On the basis of cladistics analyzes of morphological data, Phipps (1990) suggested that southwestern China and Mexico were ancestral regions of the genus and that migration of Asian and American Crataegus through the Beringian occurred. Suppose that Crataegus migrated westward from southwest China to Europe, and eastward from East Asia to North America. However, other authors have conflicting opinions. Evans and Campbell (2002) treated Crataegus as a sub-tribe of Pyrinae and, based on both molecular and non-molecular characteristics, suggested that Crataegus originated in North America. Lo et al. (2009) used sequences for the intron spacing region, chloroplast DNA regions, and LEAFY intron2 to infer species relationships from eastern Asia, western North America, eastern North America, and Europe; Their findings indicated that eastern North America and Europe may be the most recent co-origin regions of Crataegus.

China is the center of Crataegus cultivation, and the place of origin of both cultivated Crataegus species and some wild species. Based on the morphological characteristics, some researchers have suggested that 18 species and six species of Crataegus are widely distributed throughout China (Zhao and Feng, 1996; Xin and Zhang, 1997), other authors identify 20 species of Chinese Crataegus, with seven species (Dong and Li, 2015). Among these species are C. hupehensis and C. pinnatifida var. major, C. bretschneideri, and C. scabrifolia. Previous efforts to understand the evolutionary history and biogeography of Crataegus have included representatives of the Chinese Crataegus. Phipps (1990) suggested that C. scabrifolia evolved into European Crataegus and other Chinese Crataegus species (C. pinnatifida, C. hupehensis and C. sanguineae). Based on sequencing data from 14 plastid sites, Zarrei et al. (2015) treated C. maximowiczii as the division Sanguineae and suggested that the origin of the division includes east-west migration across the Beringian from western North America to eastern Asia. Lo et al. (2009) suggested that the ancestors of C. hupehensis, C. Songorica and C. pinnatifida may have spread from Europe to Asia. However, there is a lack of consensus on the direction of Chinese and European Crataegus migration.

Recent studies have examined the indeterminate (Wu et al., 2008; Zhang et al., 2008; Sheng et al., 2017) and interspecific (Su et al., 2015; Ma and Lu, 2016) relationships of Chinese Crataegus. Previous investigations used analyzes of limited morphological and molecular data, but no study explored the origin and evolution of cultivated Crataegus and related species belonging to China at the genetic level. Plant DNA contains abundant genetic information, and an increasing number of researchers have explored the relationships between species and plant diversification using molecular marker information. Molecular markers are used to identify genetic relationships within plant populations with nearly 100% reliability (Güney et al., 2018), random amplified repeats (Erfani-Moghadam et al., 2016; Zarei et al., 2017), inter-simple repeat sequences. (Sheng et al., 2017; Emami et al., 2018), and SSRs (Lo et al., 2009; Khiari et al., 2015; Brown et al., 2016) have been used extensively for genetic characterization of Crataegus, and to analyze the genetic diversity between and within the slopes of Crataegus. Among these types of molecular markers, SSR markers have gained great popularity in genetic research because they are highly polymorphic, convenient, and common. Based on dual-sequencing barcode genotyping systems, locus-specific amplification fragment sequencing (SLAF-seq) was developed for single nucleotide polymorphism (SNP) detection and genotyping using low-representation library sequencing. SLAF-seq is a high-throughput, high-resolution, low-cost method with short cycles, and has been used for molecular breeding and germplasm analysis (Huang et al., 2016). SLAF-seq does not depend on reference genome sequences and is particularly useful for species that lack a pooled reference genome (Sun et al., 2013), because it is possible to perform polymorphism analysis and develop molecular markers directly from sequence data provided by SLAF-seq (Zheng). et al., 2016). Given these advantages, SLAF-seq has been used for rapid block detection of SNP markers for polymorphism analysis, system evolution, and germplasm identification (Chen et al., 2013; Zhang et al., 2013; Xu et al., 2015).

In this study, we used SLAF-seq to gain insight into the phylogenetic relationships among seven Crataegus taxa native to China, namely C. maximowiczii, C. sanguineae, C. hupehensis, C. pinnatifida and C. pinnatifida var. major, C. bretschneideri and C. scabrifolia. These varieties are widely distributed in China and cover most of the different climatic zones of the country. The taxa sampled include the four cultivated in China (C. hupehensis, C. pinnatifida var. major, C. bretschneideri and C. scabrifolia) and three distributed near the cultivated Crataegus. C. maximowiczii and C. sanguineae are usually distributed in northeastern China. Of the cultivated cultivars, C. pinnatifida var. The primate is endemic to China and has the longest history in cultivation, and C. scabrifolia is the ancestral species of Crataegus. C. pinnatifida is a species widespread throughout China. The seven taxa were selected by other researchers as representatives of Chinese Crataegus in earlier phylogenetic studies (for example, C. pinnatifida, C. hupehensis, and C. sanguineae: Phipps, 1990; C. hupehensis, C. sanguineae, C. C. pinnatifida: Lo et al., 2009; C. maximowiczii and C. sanguineae: Zarrei et al., 2015). Previous studies provided useful information that partially resolved the phylogenetic history of Crataegus cultivation in China. However, the relationships between the species and the evolution of Crataegus cultivated in China are still unclear. In this study, SLAF-seq was used to analyze the phylogenetic relationships between 53 accessions of cultivated Crataegus and three related species in China based on their SSRs and SNPs.

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In total, 53 accessions were sampled from Chinese Crataegus, which consisted of C. maximowiczii (5 accessions), C. sanguineae (4 accessions), C. hupehensis (6 accessions), and C. pinnatifida (14 accessions) , and C. . main (12 entries), C. bretschneideri (10 entries) and C. scabrifolia (2 entries). The exogroup consisted of accessions from three exotic taxa collected from abroad (C. monogyna and C. laevigata from Europe, and C. cruss-galli from North America). The 53 Chinese accessions (Table 1) were widely distributed across a range of geographic and climatic conditions and were thus considered to be representative of the diversity of Chinese hawthorn and comprise all potential introductory sources in China.

All trees sampled were maintained in the National Repository of Hawthorn Germplasm at Shenyang Agricultural University, China (Supplementary Table S1).

Fresh leaves and fruits were sampled for morphometric measurements at maturity (from September to early October). Twenty-five leaves and fruits were sampled for each strain. Measurement methods followed the technical code for the assessment of germplasm resources of hawthorn (Crataegus L.) (Dong, 2013). Twenty-one letters were measured, including leaf shape, leaf blade lobes, leaf blade margin, and leaf color. The characteristics of qualitative traits and classifications are summarized in Supplementary Table S2.

Young leaf samples were collected, labeled, frozen in liquid nitrogen, and stored at −80 °C until DNA extraction. One gram of frozen leaf material was ground to extract genomic DNA with cetyltrimethylammonium bromide according to the Doyle and Doyle (1990) protocol. DNA quality was checked using a Nanodrop-2000 spectrophotometer (Thermo Fisher Scientific, Wilmington, DE, USA).

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Fifty-six samples were analyzed using a nuclear SSR