Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 76(3): 19-27, 2023 Firenze University Press www.fupress.com/caryologia ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-2203 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Citation: Hosseini, S., & Yaghoobi, H. (2023). Chromosome counts and kar- yotype features of different ecotypes of Allium L. species (Amaryllidaceae – Subg. Melanocrommyum) in Iran. Caryologia 76(3): 19-27. doi: 10.36253/ caryologia-2203 Received: June 27, 2023 Accepted: October 29, 2023 Published: February 29, 2024 Copyright: © 2023 Hosseini, S., & Yag- hoobi, H. This is an open access, peer-reviewed article published by Firenze University Press (http://www. fupress.com/caryologia) and distrib- uted under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, pro- vided the original author and source are credited. Data Availability Statement: All rel- evant data are within the paper and its Supporting Information files. Competing Interests: The Author(s) declare(s) no conflict of interest. Chromosome counts and karyotype features of different ecotypes of Allium L. species (Amaryllidaceae – Subg. Melanocrommyum) in Iran Shahla Hosseini*, Hiva Yaghoobi Department of Biological Science, University of Kurdistan, Sanandaj, Iran *Corresponding author. E-mail: sh.hosseini@uok.ac.ir Abstract. This study aimed to investigate the karyotypes of five Allium species, which belong to three sections of the subgenus Melanocrommyum. Bulbs from these species were collected from natural habitats in Iran, and their somatic chromosomes were analysed. The results revealed that all examined members of subg. Melanocrommyum had a basic chromosome number of x=8 and were diploid (2n=2x=16). Chromosomal data for A. saralicum and A. shatakiense are reported here for the first time. The karyo- types exhibited a variety of chromosome types and sizes, including variations observed among different accessions of the same species. In particular, A. saralicum showed satellite chromosomes ranging in size from 2.2 to 3.71 µm, located on the short arm. Seven accessions of A. saralicum, A. stipitatum, and A. haemanthoides demonstrated the presence of 1-3 B chromosomes with centromeres located in the median or sub- terminal position. Notably, the number of B chromosomes varied even among different accessions of the same species. Based on various indices, the karyotypes of the spe- cies were classified into symmetric and asymmetric groups. All karyotype asymmetry methods consistently identified A. stipitatum as the species with the most asymmet- ric chromosomes, while A. ubipetrense was recognized as the most symmetric species. This study contributes to the karyological knowledge of the genus Allium and provides valuable data for future taxonomic research. It emphasizes the significance of chromo- somal characteristics in understanding plant evolution and species diversity within the Allium genus. Keywords: Allium, B chromosome, ideogram, karyology, Melanocrommyum. INTRODUCTION Allium is a notable monocot genus comprising over 900 species, pri- marily concentrated in the eastern Mediterranean region, Southwest, and Central Asia. Within different plant communities, Allium species play a significant role (Fritsch and Abbasi, 2013). Iran exhibits diverse geograph- ic and climatic conditions, allowing Allium species to thrive in a range of habitats. These plants typically inhabit open, sunny, and relatively dry http://www.fupress.com/caryologia https://doi.org/10.36253/caryologia-2203 https://doi.org/10.36253/caryologia-2203 https://doi.org/10.36253/caryologia-2203 http://www.fupress.com/caryologia http://www.fupress.com/caryologia mailto:sh.hosseini@uok.ac.ir 20 Shahla Hosseini, Hiva Yaghoobi sites, adapting to both arid and moderately humid climates (Fritsch and Friesen, 2002). Subgenus Mel- anocrommyum of Allium encompasses Iranian species that occupy various ecological habitats, spanning from lowlands to highlands. Although they can be found in dry steppes, semi-deserts, and arid mountains, the majority of these species thrive in such environments (Fritsch and Abbasi, 2013). Currently, Iran is home to 148 recognized Allium species and subspecies, repre- senting eight subgenera and 32 sections (Fritsch and Amini Rad, 2013). Subgenus Melanocrommyum is the second-largest subgenus within Allium, encompassing approximately 170 species worldwide, classified into 20 sections (Fritsch, 2012). Chromosomes serve as the carriers of genetic infor- mation, and alterations in their number and structure play a crucial role in plant evolution (Escudero et al., 2014). Chromosomal characteristics, including number, size, and shape (karyotype), serve as defining features for numerous plant taxa across different taxonomic levels (Baltisberger and Hörandl, 2016). In the genus Allium, a basic chromosome number of x=8 is pre- dominant, although a few sections exhibit x=7, 9, and 11 (Friesen et al., 2006). Subgenus Melanocrommyum typically possesses a basic chromosome number of x=8. However, a small number of species, such as A. karat- aviense Regel and A. rnonophyllum Vved. have been identified with basic chromosome numbers of x = 9 and x=10, respectively (Fritsch and Astanova, 1998). Satellite chromosomes have proven to be a valuable cytological marker in Allium, and different types of satellites have been studied based on the centromere position and sec- ondary constrictions on chromosome arms (Dolatyari et al., 2018). Several karyological investigations have been con- ducted on Iranian Allium species, revealing symmetri- cal karyotypes composed of metacentric and submeta- centric chromosomes in the Melanocrommyum sub- genus (Pedersen and Wendelbo, 1966; Pogosian, 1983; Fritsch and Astanova, 1998; Gurushidze et al., 2010, 2012; Hosseini and Go, 2010; Akhavan et al., 2015; Dolatyari et al., 2018; Hosseini, 2018). Approximately 55% of Iranian Allium species have been karyologi- cally characterized, providing a valuable foundation for future taxonomic research (Dolatyari et al., 2018). Nonetheless, the karyological data for many species, including endemic ones in Iran, remains completely unknown. This study aims to contribute to the karyo- logical investigation of selected Iranian Allium acces- sions, expanding our knowledge of the chromosomes within the genus Allium. MATERIALS AND METHODS In this study, bulbs of five species (seven ecotypes) were collected from their natural habitats in 2018 (Fig- ure 1). The species include A. ubipetrense R.M. Fritsch, A. haemanthoides Boiss. & Reut. A. stipitatum Regel, A. saralicum R.M. Fritsch, and A. shatakiense Rech. f. Details about the collected materials can be found in Table 1. Among the species studied, A. ubipetrense was determined to be endemic to Iran, while the other spe- cies were indigenous and had distributions in various regions, including Iran, Iraq, Turkey, Tajikistan, Afghan- istan, Kazakhstan, Kyrgyzstan, Pakistan, Turkmenistan, and Uzbekistan. To break the dormancy of the bulbs, they were stored at 4 °C. Subsequently, the bulbs were rooted in wet, sterile cotton gauze in a refrigerator. For somatic chromosome analysis, fresh root tips measuring 1–1.2 cm were collected from the cultivated bulbs in the ear- ly morning. The roots underwent a pre-treatment with a-bromo naphthalene for 3 hours at 4 °C, followed by three washes with distilled water (each lasting 5 min- utes) at room temperature. The roots were then fixed overnight at 4 °C in Carnoy’s fixative, which consists of glacial acetic acid and ethanol in a 3:1 ratio. After thor- Table 1. Characterization and sampling location of the studied taxa. Section Species Species code Locality Voucher specimen Origin sect. Acanthoprason A. ubipetrense S1 Kurdistan, Marivan road, kalatarzan, jannat boo village UOK-130 Native- endemic A. haemanthoides S5 Kurdistan, Saral Area, Zardavan UOK-131 Native-non endemic sect. Melanocrommyum A. saralicum S2-E1 Kurdistan, Saral Area, hezarkanian Village UOK-113 Native-non endemic A. saralicum S2-E2 Kurdistan, Saral Area, Chatan Village UOK-114 Native-non endemic A. shatakiense S3 Kurdistan, Saral Area, zardavan UOK-126 Native-non endemic sect. Procerallium A. stipitatum S4-E1 Kurdistan, Saral Area, Kapak Village UOK-101 Native-non endemic A. stipitatum S4-E2 Kurdistan, Saral Area, zardavan UOK- 102 Native-non endemic 21Chromosome counts and karyotype features of different ecotypes of Allium L. species in Iran ough washing with distilled water, the excised roots were transferred to 70% (v/v) aqueous ethanol and stored in a refrigerator until further use. Hydrolysis was carried out by treating the root tips with 1 M HCl for 15 minutes at 60 °C. Subsequently, the root tips were stained with feulgen solution for 1 hour and then squashed in a drop of 45% (v/v) acetic acid. The best metaphase plates were photographed using a DP72 digital camera attached to the BX51 Olympus microscope. At least five metaphase plates were analyzed for each accession, and the short (S) and long arms (L) of the chromosomes were measured using IdeoKar soft- ware (http://agri.uok.ac.ir/ideokar/index.html) (Mir- zaghaderi and Marzangi 2015). The morphology of the chromosomes was determined based on the nomencla- ture proposed by Levan et al. (1964). Karyotype formu- las were established using centromere indices (CI) and arm ratio (AR). In addition to the basic karyological data such as 2n (chromosome number), x (basic chromosome number), and the total chromosome length of the haploid (HCL), several chromosomal parameters were analyzed. These parameters include the long arm (L), short arm (S), chromosome length (CL), arm ratio (AR), r value, rela- tive length of chromosome (RL), chromosome form per- centage (F %), and centromeric index (CI %). Further- more, 12 karyotype parameters were calculated to quan- tify the asymmetry of the karyotypes. These parameters Figure 1. Studied species in their habitat in saral region in Kurdistan. A, B: A. ubipetrense; C, D: A. saralicum; E, F: A. stipitatum; G, H: A. shatakiense; I; A. haemanthoides. http://agri.uok.ac.ir/ideokar/index.html 22 Shahla Hosseini, Hiva Yaghoobi include the mean centromeric asymmetry (MCA), coef- ficient of variation of chromosome length (CVCL), coef- ficient of variation of centromeric index (CVCI), total form percentage (TF %), mean centromeric index (XCI), asymmetry index (AI), degree of karyotype asymmetry (A), percentage of karyotype symmetry (S%), intra chro- mosomal asymmetry index (A1 and A2), and percentage karyotype asymmetry index (AsK %). It’s worth noting that B chromosomes were not considered in the compu- tation of these parameters due to their effects on calcu- lating asymmetry factors. RESULTS The analysis focused on metaphase plates of five spe- cies belonging to the Melanocrommyum subgenus of the genus Allium. These species are diploid, and their basic chromosome number is x=8. The chromosomal data for A. saralicum and A. shatakiense are presented for the first time in this study. Among the three species ana- lyzed (A. saralicum, A. stipitatum, A. haemanthoides), seven accessions showed the presence of 1–3 B chromo- somes with centromeres located in the median or sub- terminal position (Figure 2). Notably, the number of B chromosomes varied even among different accessions of the same species. Karyotypes of somatic complement and the ideograms of the haploid complement of studied Alliums are demonstrated in Figures 3. There were significant differences in size between the longest and shortest chromosomes in each comple- ment. Allium haemanthoides had the longest chromo- some (14.2 µm), while A. stipitatum had the shortest chromosome (5.42 µm). The mean total chromosome length (TL) ranged from 9.27 µm (A. stipitatum) to 11.37 µm (A. haemanthoides), with an overall mean value of 10.15 µm. The centromeric index (CI) of the comple- ments varied from 37% (A. stipitatum) to 41% (A. ubipe- trense). Based on the nomenclature proposed by Levan et al. (1964), two chromosome types, ‘m’ (centromere at the median region) and ‘sm’ (centromere at the submedian region), formed six different karyotype formulas (Table 2). Additionally, there were pairs of satellites with sizes ranging from 2.2 to 3.71 µm in A. saralicum (S2-E1 and S2-E2) located on the short arm. The karyotypes of all five species were classified into the 1A, 1B, and 2B classes of Stebbins classification (Stebbins 1971). Various methods were used to assess karyotype asymmetry, and most methods identified dif- ferent species as symmetric or asymmetric. For example, A. ubipetrense had the highest value of total form per- centage (TF %) at 41.56 (indicating the most symmet- ric species), while A. stipitatum had the lowest value at 37.52 (the most asymmetric species). The coefficient of variation (CV %) showed the highest value in A. stipi- tatum (26.88%, the most asymmetric) and the lowest value in A. ubipetrense (11.77%, the most symmetric). The highest XCA value (mean centromeric asymmetry) was observed in A. stipitatum (24.11%), while the low- est value was found in A. shatakiense (17.95%). The mean coefficient of variation of centromeric index (CVCI) was determined as 13.56 µm, ranging from 7.7 µm (A. ubi- petrense) to 14.32 µm (A. stipitatum). Allium stipitatum had the highest value of asymmetry index (A) at 0.24, while A. ubipetrense and A. shatakiense showed the low- est value at 0.17. The highest and lowest values of per- centage of karyotype symmetry (S %) were observed in A. ubipetrense (68.83) and A. stipitatum (43.20), respec- tively. Allium stipitatum had the highest value of per- centage karyotype asymmetry index (AsK %) at 62.24 but A. ubipetrense showed the lowest value (54.90) (Table 2). According to the data presented in Table 2, all the karyotype asymmetry methods consistently identified A. stipitatum as the species with the most asymmetric chromosomes, while A. ubipetrense was recognized as the most symmetric species. These findings highlight the distinct karyotype characteristics and asymmetry levels among the studied Allium species. DISCUSSION The karyotypic characteristics of several Allium spe- cies, including A. saralicum, A. shatakiense, A. stipitat- um, A. ubipetrense and A. haemanthoides are discussed in this study. All studied taxa were found to be diploid with a chromosome number of 2n=2x=16. Allium stipi- tatum, on the other hand, exhibited the same diploid chromosome number of 2n=2x=16, which is consistent with previous studies (Fritsch and Astanova, 1998; Ohri and Pistrick, 2001; Oroji Salmasi et al., 2019). However, reports also exist of A. stipitatum having a basic chro- mosome number of 2n=16 and 2n=48 (Pogosian 1983). In A. saralicum, a pair of satellite chromosomes was observed, located on the short arm of the chromosomes. The present study identified two chromosome types, ‘m’, ‘sm’, and ‘st’, in five different Allium species. The karyo- typic analysis, combined with previously published data on the Melanocrommyum subgenus, indicated a com- mon symmetric karyotype with 2-8 metacentric and 0-4 submetacentric chromosomes, as well as 0-4 subtelocen- tric chromosome pairs. Chromosomes in the Acanthoprason section (A. ubi- petrense and A. haemanthoides) displayed a median or 23Chromosome counts and karyotype features of different ecotypes of Allium L. species in Iran submedian centromere position and a gradual decline in arm lengths. Allium ubipetrense had chromosomes with centromeres only in the median position, while A. hae- manthoides exhibited submedian centromeres and one B Figure 2. Mitotic metaphase plates of the investigated accessions of selected Allium, subg. Melanocrommyum: A, B: A. ubipetrense; C-F: A. sarali- cum (E1); G, H; A. saralicum (E2); I-K; A. stipitatum (E1); L; A. stipitatum (E2); M; A. haemanthoides; N,O; A. shatakiense. Scale bars = 10µm. 24 Shahla Hosseini, Hiva Yaghoobi chromosome in addition to metacentric ones. The coef- ficient of variation for centromere index (CVCI) in A. stipitatum was more than twice (14.32) that of A. ubipe- trense (6.35). The coefficient of variation for centromere length (CVCL) index confirmed A. ubipetrense as having the most symmetric chromosomes. A study by Dolatyari et al. (2018) on A. haemanthoides did not detect any B chromosomes but identified two satellite (SAT) chromo- somes with centromeres positioned between the median and sub median positions. Allium ubipetrense is primarily found in the north- western parts of the Zagros mountain range in Iran. This species exhibits different morphotypes, which sometimes leads to misidentifications. Allium haeman- Figure 3. Idiograms and karyotypes of the investigated accessions of selected Allium subg. Melanocrommyum. A: A. ubipetrense; B; A. sarali- cum; C: A. shatakiense; D: A. stipitatum; E: A. haemanthoides. Scale bars = 10µm. 25Chromosome counts and karyotype features of different ecotypes of Allium L. species in Iran thoides, classified under sect. Acanthoprason, shares one subgroup with A. ubipetrense and A. kurdistani- cum from the Kurdistan province, based on molecular markers (ITS sequences of nuclear rDNA) (Fritsch and Abbasi 2013). Chromosomal parameters suggest a high probability of differentiation between the two species, A. haemanthoides and A. ubipetrense, into different subgroups. Regarding the Melanocrommyum section, A. sarali- cum and A. shatakiense demonstrated closely similar values for all parameters and indices. The karyotypes of both species consisted of metacentric and sub meta- centric chromosomes, with one B chromosome and cen- tromeres in the median position. This study reports the symmetric karyotype of section Melanocrommyum for the first time in A. saralicum and A. shatakiense. Previ- ous studies have also shown this symmetric karyotype in the Melanocrommyum section (Fritsch & Astanova 1998; Hosseini and Go, 2010; Akhavan & al. 2015; Dola- tyari et al., 2018; Hosseini, 2018). Allium stipitatum, the only species analyzed karyo- logically in section Procerallium, was also diploid with 2n=16. However, its karyotype differed from the other species in subg. Melanocrommyum, as it included 1-3 subtelocentric B chromosomes. Among the studied taxa, Allium stipitatum exhibited the highest values for all asymmetric indices, such as CVCL (26.88), CVCI (14.32), ASK% (62.24), and the lowest values for TF% (37.52), rvalue (0.6), F% (4.6), and CI (0.37). Nevertheless, when it comes to distinguishing closely related taxa at the section level, the uniformity of karyotypes and comparable chromosome counts do not hold significant value. This research focused on a five species across three different sections, and it would be inaccurate to apply these findings universally to the entire Allium genus. To achieve a clearer understanding Table 2. Mean chromosomal and karyotypic parameters of Allium spp. S1: A. ubipetrense S2: A. saralicum, S3: A. shatakiense, S4: A. stipi- tatum, S5: A. haemanthoides. Parameters Species Mean Range S1 S2 S3 S4 S5 Min Max S (µm) 4.09 4.30 3.89 3.47 4.52 4.05 3.47-4.52 S4 S5 L (µm) 5.79 6.55 5.50 5.79 6.84 6.09 5.50-6.84 S3 S5 TL (µm) 9.89 10.86 9.40 9.27 11.37 10.15 9.27-11.37 S4 S5 AR 1.44 1.63 1.45 1.69 1.54 1.55 1.44-1.69 S1 S4 r value 0.69 0.64 0.70 0.60 0.66 0.65 0.60-0.70 S4 S3 HCL 79.21 89.03 75.27 75.74 93.61 82.57 75.27-93.61 S3 S5 RL% 12.49 12.24 12.49 12.25 12.14 12.32 12.14-12.49 S5 S1,S3 F % 5.15 4.86 5.13 4.60 4.83 4.91 4.60-5.15 S4 S1 CI 0.41 0.38 0.40 0.37 0.39 0.39 0.37-0.41 S4 S1 TF % 41.56 39.65 41.20 37.52 40.00 39.98 37.52-41.56 S4 S1 CVCL 11.77 19.89 20.72 26.88 18.90 19.63 11.77-26.88 S1 S4 A1 0.30 0.33 0.28 0.37 0.32 0.32 0.28-0.37 S3 S4 A2 0.11 0.19 0.20 0.26 0.18 0.18 0.11-0.26 S1 S4 AI 185.24 155.31 170.48 214.10 174.87 180 155.31-214.10 S2 S4 AsK % 54.90 60.34 58.43 62.24 58.79 58.94 54.90-62.24 S1 S4 S% 68.83 52.49 51.91 43.20 54.90 54.26 43.20-68.83 S4 S1 A 0.17 0.21 0.17 0.24 0.19 0.19 0.17-0.24 S1,S3 S4 XcA 17.95 21.72 17.53 24.11 19.74 20.21 17.53-24.11 S3 S4 XcI 0.17 0.39 0.41 0.37 0.40 0.34 0.17-0.41 S1 S3 CVCI 6.35 12.47 12.09 14.32 11.45 13.56 6.35-14.32 S1 S4 Species S1 S2-E1 S2-E2 S3 S4-E1 S4-E2 S5 STa 1A 1A 1A 1A 2B 2B 1B KFb 16m 10m+6sm 10m+6sm+ 1smB 12m+4sm+1smB 8m+8sm+1stB 12m+4sm+3stB 10m+6sm+1smB a ST Stebbin’s (1971) classification; b KF Karyotype formula. 26 Shahla Hosseini, Hiva Yaghoobi of sectional boundaries, it would be beneficial to con- duct future studies that involve a larger variety of spe- cies from diverse sections, while also examining chro- mosomal karyotypes in greater detail. CONCLUSION In conclusion, the karyological investigation of five Allium species belonging to the Melanocrommyum sub- genus provides valuable insights into the chromosomal characteristics and karyotype diversity within this genus. The species studied, A. ubipetrense, A. haemanthoides, A. stipitatum, A. saralicum, and A. shatakiense, exhibited diploid chromosome numbers and a basic chromosome number of x=8. However, some variations were observed, such as the presence of 1-3 B chromosomes in certain accessions. The karyotypes of these species displayed a combination of metacentric, submetacentric, and subte- locentric chromosomes, with centromeres positioned at the median or submedian regions. Additionally, satellite chromosomes were observed in A. saralicum. The karyo- type asymmetry analysis revealed variations among the species, with A. stipitatum exhibiting the most asymmet- ric chromosomes and A. ubipetrense displaying the most symmetric chromosomes. These findings contribute to the understanding of the genetic diversity and evolutionary patterns within the Allium genus, particularly the Melanocrommyum subgenus. The data obtained from this study adds to the existing knowledge of Allium species in Iran and serves as a foundation for future taxonomic and evolutionary research. Further investigations of karyological charac- teristics and chromosomal variations in other Iranian Allium species would provide a more comprehensive understanding of their genetic makeup and phylogenetic relationships. ACKNOWLEDGMENTS This research was funded by the University of Kurd- istan (UOK), Iran, as a research grant (11.99.5722). REFERENCES Akhavan, A., Saeidi, H., Zarre, S., & Rahiminejad, M. (2015). Chromosome Numbers and Karyotype Fea- tures of Selected Species of Allium L. (Amaryllidace- ae) Sect. Acanthoprason in Iran. Iranian Jornal of Botany, 21(2), 158-164. Baltisberger, M., & Hörandl, E. (2016). 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