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Download Download Firenze University Press Caryologia www.fupress.com/caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Cytogenetic and molecular studies of the Egyptian Capsella bursa-pastoris (Brassicaceae) Citation: W.M. Amer, R.A. Hassan, A.S. Abdo (2020) Cytogenetic and molecu- lar studies of the Egyptian Capsella bursa-pastoris (Brassicaceae). Caryo- Wafaa M. Amer, Rania A. Hassan*, Amany S. Abdo logia 73(3): 45-54. doi: 10.13128/caryolo- gia-867 Botany and Microbiology Department, Science Faculty, Cairo University, Egypt * Corresponding authors. E-mail: [email protected]; [email protected] Received: February 23, 2020 Accepted: April 19, 2020 Abstract. Capsella bursa-pastoris (Brassicaceae) is one of the most successful tetra- ploid species in the world. It showed high morphological diversity within Egyptian Published: December 31, 2020 populations. Morphological investigations of herbarium specimens and fresh col- Copyright: © 2020 W.M. Amer, R.A. lected populations grouped them under three distinguished morphotypes (Lobed Hassan, A.S. Abdo. This is an open “L”; Simple “S” and Lobed-Simple “LS”) depending mainly on the basal leaves struc- access, peer-reviewed article pub- ture. This high degree of phenotypic variation has received our critical attention. Until lished by Firenze University Press recently, the previous studies on C. bursa-pastoris attributed its phenotypic variation (http://www.fupress.com/caryologia) to environmental factors. But in Egypt, these three morphotypes were traced in mixed and distributed under the terms of the populations along with the species geographical range, so the environmental factors Creative Commons Attribution License, have no influence on their distribution or phenotypic variation. Accordingly, our pri- which permits unrestricted use, distri- mary concern in this study was to determine the factors controlling this variation. bution, and reproduction in any medi- um, provided the original author and The cytogenetic studies revealed that the three identified morphotypes are three dis- source are credited. tinct genotypes with three different chromosome numbers: 2n=2x=16 (diploid) for “L”; 2n=3x=24 (triploid) for “S”; 2n=4x=32 (tetraploid) for “LS”. The triploid genotype “S” Data Availability Statement: All rel- showed rare occurrence among the studied populations and is postulated to be a new evant data are within the paper and its record of a hybrid in Egypt. Karyotyping of the three genotypes showed significant Supporting Information files. differences in the genome and chromosomes relative lengths. Molecular study using Competing Interests: The Author(s) cpSSR technique supported the cytogenetic results and differentiated the three studied declare(s) no conflict of interest. genotypes. The retrieved results revealed that the phenotypic diversity within the Egyp- tian C. bursa-pastoris populations is genetically controlled. Keywords: Capsella bursa-pastoris, cytogenetic studies, genotypes, karyotyping, molecular study, phenoplasticity. INTRODUCTION Brassicaceae (Cruciferae) or mustard family is one of the largest Angio- sperm families, it comprises 3977 species and 341 genera and 52 tribes (Kief- er et al., 2014). One of the most important genera in the Brassicaceae is genus Capsella Medik. Molecular systematic studies confirmed that genusCapsella belongs to the tribe Camelineae (Neuffer et al., 2014). This genus is an excel- lent model for molecular evolutionary studies due to its phylogenetic rela- tions within the Brassicaceae. Studying its genetics, speciation and sympatric distribution is important for agricultural matters. Genus Capsella is represented worldwide by five species: the two self- compatible tetraploid C. bursa-pastoris (L.) Medik and C. thracica Velen Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 73(3): 45-54, 2020 ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.13128/caryologia-867 46 Wafaa M. Amer, Rania A. Hassan, Amany S. Abdo (2n=4x=32); the self-incompatible diploid C. grandiflora MATERIALS AND METHODS (Fauche & Chaub) Boiss; the two self-compatible diploid C. rubella Rent. and C. orientalis Klokov (Hurka et al., Morphological study 2012; Neuffer et al., 2014). These species differ greatly in their geographical distribution, where, C. grandiflora is The morphological investigations of C. bursa-pasto- limited to northwestern Greece and Albania; C. rubella ris were carried on 36 old populations deposited as her- has a broad Mediterranean / central European distribu- barium specimens in Cairo University Herbarium (CAI) tion; C. orientalis is found from Eastern Europe to Cen- and Assiut University Herbarium (ASTU). In addition tral Asia (Hurka et al., 2012); C. thracica is endemic to to 66 fresh populations collected from Menoufia (Abu Bulgaria (Neuffer et al., 2014). Sleem village), Faiyum (Sinnuris district, El Siliene) and Capsella bursa-pastoris (Shepherd’s Purse) is an El Saff regions during our field work conducted in 2016- annual to biennial species, extremely variable in size and 2018. The studied specimens (old & fresh) from differ- leaf form, distinguished by terminal and axillary raceme ent distribution localities are shown in Table 1. From inflorescences. Its silicula fruit is obcordate-obtriangular 66 fresh populations, 25 specimens/ population were in shape (Amer et al., 2019). This species is the second undergone morphological investigations using different most common flowering plant in the world (Zhou et al., morphological criteria of leaves, inflorescence and fruit. 2001), grows as a common weed of agriculture in almost Acronyms of herbaria follow Thiers (2019). all countries of the world from tropical to subarctic hab- The morphological investigations distinguished itats (Holm et al., 1979), and shows a high phenotypic three morphotypes of C. bursa-pastoris in all the stud- plasticity (Korsmo, 1954; Holzner and Numata, 1982). ied populations namely: (L) Lobed, (S) Simple and (LS) Accordingly, the evolution of polyploidy and weediness Lobed-Simple (Amer et al., 2019). in C. bursa pastoris is interesting to agricultural research (St Onge, 2010). Cytogenetic studies Many taxonomic studies were carried out on this species depending on morphological characters and Sample preparation resulted into many species, subspecies, varieties, micro- species, biotypes, and segregates (Aksoy et al., 1998; For cytogenetic studies, the specimens collected Aksoy et al., 1999; Neuffer, 2011). from Faiyum region (marked with * in Table 1) were A considerable amount of literature has attributed selected to nullify the environmental factors. Seeds of the phenotypic variation in C. bursa-pastoris to environ- 30 specimens representing the three morphotypes (10/ mental or geographical factors like seasonality, tempera- morphotype) were collected, soaked in distilled water ture, shade, rainfall, latitudinal and altitudinal gradients for 2 hours, and germinated at room temperature. Root (Almquist, 1929; Neuffer, 1989; Neuffer and Bartelheim, tips of about 1 cm length were treated with colchicine 1989; Stace, 1989; Neuffer, 1990; Aksoy, 1996; Aksoy et (C22H25NO6, 0.025 %) for 2 hours at room temperature, al., 1999; Neuffer and Hoffrogge 2000). and washed thoroughly with distilled water. Fixation In Egypt, Capsella is a monospecific genus repre- was done using ethyl alcohol: glacial acetic acid (3:1, v/v). sented by C. bursa-pastoris (Boulos, 1999). The field Samples were washed thoroughly with water and hydro- study and morphological investigations of this species lyzed using 1 N HCl at 64° C for 5 minutes. The slides – based on the leaves, inflorescence and fruit charac- were prepared by squashing the root tips using 45% ace- ters – showed the presence of high degree of phenotypic tic acid and stained with aceto orcein solution. variation and revealed the presence of three morpho- types namely: Lobed “L”, Simple “S” and Lobed-Simple “LS” (Amer et al., 2019). These three morphotypes were Chromosome count and karyotyping traced in mixed populations along with the species geo- graphical range, so the environmental factors have no Chromosome count was performed on mitotic meta- influence on their distribution or phenotypic variation. phase cells. For each morphotype, ten clearly observable Therefore, this study aims to determine the factors metaphase cells from ten individuals were selected and controlling the phenotypic variation of C. bursa-pastoris photographed using standard and high resolution auto- morphotypes through cytogenetic and molecular studies mated karyotyping software processing (Leica CW4000). to clarify the impact of genetic diversity on their pheno- Metaphase chromosomes of each morphotype were plasticity. placed in pairs, arranged and numbered in order of size, with keeping in view the centromere position to consti- Cytogenetic and molecular studies of the Egyptian Capsella bursa-pastoris (Brassicaceae) 47 tute a karyotype. The length of the short arm (p) and the chosen for molecular study (4 Lobed, 4 Simple and 6 long arm (q) was measured for each chromosome, and Lobed-Simple). the total length (TL=p+q) was calculated. The relative length (RL) of the chromosomes (TL / sum TL × 100) and the mean relative length (MRL) of each chromosome DNA extraction pair were calculated. The centromeric index (CI) was estimated by (P / TL x 100), the mean centromeric index A total genomic DNA was extracted from 1 g young (MCI) was calculated to represent the centromeric index leaves using CTAB
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