Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 76(2): 59-65, 2023 Firenze University Press www.fupress.com/caryologia ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-2304 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Citation: R.N. Chougule, P.V. Desh- mukh, P.E. Shelke, V.J. Patil, M.M. Lekhak (2023). Cytogenetical studies of some Convolvulaceae members from the Western Ghats, India reveal uni- formity in karyotypes. Caryologia 76(2): 59-65. doi: 10.36253/caryologia-2304 Received: September 8, 2023 Accepted: October 29, 2023 Published: December 31, 2023 Copyright: © 2023 R.N. Chougule, P.V. Deshmukh, P.E. Shelke, V.J. Patil, M.M. Lekhak. 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. Cytogenetical studies of some Convolvulaceae members from the Western Ghats, India reveal uniformity in karyotypes R.N. Chougule, P.V. Deshmukh, P.E. Shelke, V.J. Patil, M.M. Lekhak* Angiosperm Taxonomy Laboratory, Department of Botany, Shivaji University, Kolhapur, Maharashtra 416004, India *Corresponding author. E-mail: mml_botany@unishivaji.ac.in Abstract. Karyotypes of six Ipomoea and one Merremia species were studied. Chromo- some counts of 2n = 30 for I. corymbosa, I. kotschyana, I. marginata f. candida and M. rhyncorhiza) and 2n = 60 for I. ochracea were observed for the first time. Meiotic course of five Ipomoea species was also examined for the first time and two counts, i.e. n = 15 and n = 30 were recorded. Meiosis was found normal. Karyotypes of the stud- ied species exhibited uniformity. All the species had chromosomes with median region centromeres and karyotypes were symmetrical (Stebbins’ 4A category). I. marginata f. candida had the shortest chromosomes with mean length of 0.99 ± 0.02 µm. I. ochra- cea and I. corymbosa had the longest chromosomes with mean length of 1.22 ± 0.05 µm and 1.22 ± 0.01 µm, respectively. As chromosomes were small and exhibited uni- form morphology, fluorescent banding or fluorescent in-situ hybridization can be use- ful to differentiate the karyotypes and understand species interrelationships. Keywords: chromosomes, morning glory, karyotype symmetry, Western Ghats. INTRODUCTION Convolvulaceae, called as bindweeds or morning glories are a family of about 58 genera and more than 1977 species worldwide (POWO 2023). In India, there are about 24 genera that include about 164 species of which 27 are endemic (Dash and Mao 2020; Singh et al. 2015). These species are most- ly herbaceous to woody vines, rarely herbs, shrubs, or trees. India is home to a rich diversity of Convolvulaceae due to the presence of diverse habitats and exposure to introduced species. Ipomoea L. is a large genus that comprises more than 800 species distributed over tropical and subtropical regions of the world (Wood et al. 2020). In India, the genus is represented by (64 taxa) 55 species and 9 infraspecific taxa, of which only three species, I. clarkei Hook.f, I. laxiflora H.J.Chowdhery & Debta and I. salsettensis Santapau & Patel are endemic (Dash and Mao 2020; Singh et al. 2015). Some taxa such as I. cairica var. semine-glabro (Blatt. & Hallb.) Bhandari, I. marginata f. can- dida (Naik & Zate) Das Das & Lakshmin., I. deccana var. lobata (C.B.Clarke) 60 R.N. Chougule et al. S.C.Johri, I. nil var. himalaica (C.B.Clarke) S.C.Johri, I. obscura f. concolor Naik & Zate, I. pes-caprae var. perunkulamensis P.Umam. & P.Daniel have also been reported as endemic to India (Kattee 2019). The West- ern Ghats consists of 42 taxa (37 species, 03 subspecies and 02 varieties) (Lekhak et al. 2018). Merremia Dennst. ex Endl., on the other hand includes 49 species distrib- uted over tropical and subtropical regions of the world (POWO 2023). In India, Merremia comprises 17 species, of which two species, M. rajasthanensis Bhandari and M. rhyncorhiza (Dalzell) Hallier f. are endemic to the coun- try (Dash and Mao 2020; Singh et al. 2015). Convolvulaceae members are economically very important. For instance, I. batatas (L.) Lam., popularly known as sweet potato, is a rich source of energy for humans as well as animals. The plant parts of I. pes- caprae (L.) R.Br. have been traditionally used to treat gastrointestinal-related disorders and symptoms, such as dysentery, ulcer, abdominal pain, cramps and stom- ach aches (Emendörfer et al. 2005; Pereda-Miranda et al. 2005). Many members such as Convolvulus L., Ipo- moea, Stictocardia Hallier f., etc. are used as ornamen- tals. Moreover, the members of the family serve as an important source of ergoline alkaloids (Fig. 1) that have been used to treat nervous system disorders like con- vulsions, epilepsy, migraine, Parkinson’s disease or are used in childbirth and weaning (Groeger and Floss 1998; Mutschler et al. 2001; Schardl et al. 2006; Chen et al. 2018). On account of its tremendous utility in medicinal and ornamental fields, the family is cytogenetically well explored. In India, somatic chromosomes of these mem- bers have been the focal point for the majority of studies (Vij et al. 1974, 1977; Bir and Sidhu 1975; Bir et al. 1978; Roy 1979; Sampathkumar 1979; Rao and Mwasumbi 1981; Sinha and Sharma 1992; Rane et al. 2012; Lekhak et al. 2018; Lawand et al. 2019; Ramanpreet and Gup- ta 2018). Most of these studies recorded chromosome counts for species of Ipomoea, Argyreia Lour., Merremia and Operculina Silva Manso. The most common diploid chromosome number reported in these genera is 2n = 30. Although, 2n with 18, 22, 28, 32, 38, 58 and 60 chro- mosomes have also been reported in some species (Yeh and Tsai 1995; Lekhak et al. 2018; Lawand et al. 2019). Instances of polyploidy are usually rare. Nevertheless, polyploidy has been reported in Ipomoea batatas which is a hexaploid (2n = 6x = 90) (Sinha and Sharma 1992; Vij et al. 1977). The present investigation aims at generating new cytogenetical data for the f lowering plants (wild or introduced) of the Western Ghats, India. Since Convol- vulaceae members are of horticultural and medicinal importance, chromosomal information would be useful in understanding their genetic potential, phylogenetic relationships and breeding strategies. Accordingly, here we provide comparative karyotypes of seven species. Meiotic chromosomes were studied for five Ipomoea species. Karyological analysis of taxa was based on kar- yological parameters such as diploid chromosome num- ber (2n), mean chromosome length (MCL), total haploid chromosome length (THL), mean centromeric asym- metry (MCA) and coefficient of variation of chromosome length (CVCL). MATERIAL AND METHODS The plant materials for the present study were col- lected from different localities in Gujarat and Maharash- tra state. Plants were cultivated in Lead Botanical Gar- den, Department of Botany, Shivaji University, Kolhapur and the voucher specimens deposited in the Herbarium of Shivaji University, Kolhapur (SUK) (Table 1). For mitotic preparations, seeds were nicked near the hilum with the help of a sharp razor, and then placed in the petri dish on a wet blotting paper. Well-grown root tips (1.5-3 cm long) were harvested from the germinating seeds and pre-treated with a saturated solution of para- Dicholorobenzene (pDB) for 4-5 h at 9±3oC. Further, the root tips were hydrolysed in 1N HCl and squashed in 2% propionic orcein. For meiotic studies, appropri- ately sized flower buds were fixed in Carnoy’s solution and smears of anthers from floral buds were stained using 2% propionic orcein. Suitable somatic and meiotic plates from freshly prepared slides were photographed with Leica DM 750 microscope with attached camera at 1000X magnification. Five cells with well-spread meta- phase chromosomes were selected for karyotype analy- sis. Nomenclature of chromosomes follows Levan et al. (1964). Karyotype asymmetry was ascertained using CVCL (coefficient of variation of chromosome length) Figure 1. Structure of Ergoline alkaloid. 61Cytogenetical studies of some Convolvulaceae members from the Western Ghats, India reveal uniformity in karyotypes and MCA (mean centromeric asymmetry) as suggested by Peruzzi and Eroğlu (2013). RESULTS In the present investigation mitotic metaphase chro- mosomes of seven species were studied. Meiotic study was performed on five species. Ipomoea ochracea (Lindl.) Sweet exhibited 2n = 60 chromosomes while the rest of the species had 2n = 30 chromosomes. Comparative kar- yotypes of all the species investigated are summarized in Table 2. Fig. 2 illustrates the mitotic metaphases while Fig. 3 depicts ideogram. Chromosomes were with medi- an region centromeres, and hence the karyotype formula 15m (I. corymbosa, I. kotshyana, I. laxiflora, I. margina- ta f. candida and M. rhyncorhiza) or 30m (I. ochracea). Four B-chromosomes were observed in I. salsettensis (Fig. 2f) and the karyotype formula was 15m+4B. The highest mean chromosome length (MCL) (1.22 ± 0.05 and 1.22 ± 0.01 µm) was recorded in the case of I. cor- ymbosa and I. ochracea whereas the lowest (0.98 ± 0.03 µm) in I. kotschyana (Table 2). Total haploid chromo- some length (THL) ranged from 14.65 µm (I. kotschya- na) to 36.61 µm (I. ochracea). I. corymbosa showed maxi- mum value (1.74) for R (ratio of largest and the smallest chromosome of the complement) while the minimum (1.58) was recorded for I. laxiflora. MCA was found to be the lowest (6.85) for M. rhyncorhiza and the highest (14.65) for I. kotshyana. The lowest CVCL was recorded for I. marginata f. candida (12.28) and the highest for I. corymbosa (14.70) (Table 2). Meiotic studies were carried out in five species (I. horsfalliae, I. laxiflora, I. marginata f. candida, I. ochracea, I. salsettensis). Meiosis was found to be nor- mal. Pollen mother cells (PMCs) of I. ochracea showed 30 bivalents (n = 30) at diakinesis (Fig. 2k) whereas rest of the species (I. horsfalliae, I. laxiflora, I. marginata f. candida and I. salsettensis) showed 15 bivalents (n = 15) (Fig. 2h, i, j, l). Table 1. Collection localities and voucher details of studied species. Sr. No. Taxa Collection locality Voucher specimen 1. Ipomoea corymbosa (L.) Roth Shivaji University Campus, Kolhapur, Maharashtra RNC 127 2. I. horsfalliae Hook. Uajalaiwadi, Kolhapur district, Maharashtra RNC 121 3. I. kotschyana Hochst. ex Choisy Dinodhar hill, Nakhatrana taluka, Kutch district, Gujarat RNC 206 4. *I. laxiflora H.J.Chowdhery & Debta Gujarat RNC 277 5. *I. marginata f. candida (Naik & Zate) Das Das & Lakshmin. Shivaji University Campus, Kolhapur, Maharashtra RNC 207 6. I. ochracea (Lindl.) Sweet Shivaji University Campus, Kolhapur, Maharashtra RNC 120 7. *I. salsettensis Santapau & Patel Rajapur, Ratnagiri district, Maharashtra RNC 278 8. *Merremia rhyncorhiza (Dalzell) Hallier f. Chaukul, Sawantwadi taluka, Sindhudurg district, Maharashtra RNC 217 *Indicates endemic species. Table 2. Comparative karyotypes. Sr. No. Taxa 2n Range of chromosome length ± SE (µm) Arm ratio (r) ± SE THL (µm) MCL ± SE (µm) MCA CVCL R St Haploid karyotype formulae 1. I. corymbosa 30 1.56 ± 0.02 - 0.90 ± 0.03 1.16 ± 0.01 18.35 1.22 ± 0.05 7.42 14.70 1.74 4A 15m 2. I. kotschyana 30 1.21 ± 0.01 - 0.74 ± 0.01 1.22 ± 0.01 14.65 0.98 ± 0.03 14.65 13.16 1.65 4A 15m 3. I. laxiflora 30 1.35 ± 0.04 - 0.86 ± 0.02 1.15 ± 0.04 15.76 1.05 ± 0.02 7.05 12.66 1.58 4A 15m 4. I. marginata f. candida 30 1.20 ± 0.01 - 0.74 ± 0.02 1.22 ± 0.02 14.80 0.99 ± 0.02 8.84 12.28 1.65 4A 15m 5. I. ochracea 60 1.57 ± 0.02 - 0.94 ± 0.01 1.21 ± 0.03 36.61 1.22 ± 0.01 9.26 12.85 1.66 4A 30m 6. I. salsettensis 30 1.51 ± 0.07 - 0.95 ± 0.04 1.16 ± 0.04 17.85 1.19 ± 0.04 7.39 12.68 1.59 4A 15m + (0–4B) 7. M. rhyncorhiza 30 1.38 ± 0.04 - 0.85 ± 0.02 1.15 ± 0.01 16.31 1.09 ± 0.04 6.85 12.83 1.61 4A 15m THL = Total haploid length, MCL = Mean chromosome length, MCA = Mean centromeric asymmetry, CVCL = Coefficient of variation of chromosome length, R = ratio of the longest to shortest chromosome of a complement, St = karyotype asymmetry. 62 R.N. Chougule et al. DISCUSSION According to Löve and Löve (in Sinha and Sharma 1992) x = 5 is the primary basic chromosome number for the family Convolvulaceae while x = 14 and x = 15 are secondarily derived basic numbers. Darlington and Wylie (1955) and Vij et al. (1977) considered x = 14 and x = 15 as the basic chromosome numbers for the genus Ipomoea and Merremia. Most of the Indian species exhibit a diploid chromosome number of 2n = 30. In the present studies we observed 2n = 30 chromosomes in six species whereas 2n = 60 was found in I. ochracea. Earli- er, the count of 2n = 60 has been observed as an instance of tetraploidy in I. wightii (Wall.) Choisy, I. plebeia R.Br., I. lonchophylla J.M.Black, I. racemigera F.Muell. & Tate, I. ramonii Choisy, I. tiliacea (Willd.) Choisy and I. arbo- rescens (Humb. & Bonpl. ex Willd.) G.Don (Jones 1964; Yen et al. 1992; Kattee 2019) whereas, I. batatas shows hexaploidy, i.e. 2n = 90 chromosomes (Sinha and Shar- ma 1992; Pitrez et al. 2008). Some species such as I. staphylina Roem. & Schult., I. purpurea (L.) Roth pos- sess 2n = 32 chromosomes (Roy 1979; Sampathkumar 1979; Dhar 2015) whereas I. coptica Verdc., I. diversifo- lia R.Br., I. pes-tigridis L. show 2n = 28 chromosomes (Sampathkumar 1979; Lekhak et al. 2018; Bir and Sidhu 1975). I. triloba L. is reported to have 2n = 38 chromo- somes (Wang et al. 1998). These chromosome numbers not confirming to base numbers x = 14 and x = 15 can be attributed to polyploidy, aneuploidy or dysploidy or to the occurrence of polysomaty (Vij et al. 1977; Sampath- kumar 1979; Lekhak et al. 2018). Sampathkumar (1979) studied the karyomorphology of eighteen Ipomoea species whereas Rane et al. (2012) studied ten species. In both the studies chromosomes Figure 2. Mitotic metaphase and meiotic chromosomes: (a) Ipo- moea corymbosa (2n = 30); (b) I. kotschyana (2n = 30); (c) I. laxi- flora (2n = 30); (d) I. marginata f. candida (2n = 30); (e) I. ochracea (2n = 60); (f) I. salsettensis (2n = 30+4B) Arrowheads show B-chro- mosomes; (g) Merremia rhyncorhiza (2n = 30). (h-l) PMCs at diaki- nesis: (h) I. horsefalliae (n = 15); (i) I. laxiflora (n =15); (j) I. margi- nata f. candida (n = 15); (k) I. ochracea (n = 30); (l) I. salsettensis (n = 15). Scale bars = 5 μm. Figure 3. Ideograms of Ipomoea and Merremia species: (a) I. corym- bosa; (b) I. kotschyana; (c) I. laxiflora; (d) I. marginata f. candida; (e) I. salsettensis; (f) I. ochracea; (g) M. rhyncorhiza. Scale bars = 1 μm. 63Cytogenetical studies of some Convolvulaceae members from the Western Ghats, India reveal uniformity in karyotypes with median and submedian centromeres were observed. Nakajima (1963) also reported median, submedian and terminal centromere in I. lacunosa L. and I. violacea L. and hence, the karyotype was considered asymmetrical. In the present study we observed all the chromosomes with median region centromere and karyotype was highly symmetrical (Table 2). Similar observations have been made for I. clarkei and I. diversifolia (Lekhak et al. 2018) and in some NE Brazilian Ipomoea species (Pitrez et al. 2008). Kattee (2019) reported two counts, i.e. 2n = 30 (I. laxiflora, I. salsettensis and I. tenuipes Verdc.) and 2n = 60 (I. wightii). She observed chromosomes with median region centromere. In the present study, the chromosome number for I. laxiflora and I. salsettensis have been confirmed, although four B-chromosomes were observed in I. salsettensis for the first time (Fig. 2f). B-chromosomes (0-1) have also been reported in other Ipomoea species such as I. mutabilis Lindl. and I. palmata Forssk. (white flowered type) by Vij et al. (1977) whereas Yen and Tsai (1995) observed B-chromosomes (0-3) in some Ipomoea species from Taiwan. Sampathku- mar (1979) recorded the presence of a satellite chromo- some pair and secondarily constricted chromosomes in Ipomoea species. Similarly, Pitrez et al. (2008) observed satellite chromosomes in some Ipomoea species from NE Brazil inselbergs. In the present investigation, we could not find satellite chromosomes or secondarily constrict- ed chromosomes. Chromosome length ranged from 2.13 to 4.79 µm in I. carnea Jacq. and 1.25 to 2.67 µm in I. aquatica For- ssk. (Rane et al. 2012). Lekhak et al. (2018) reported the shortest chromosome in I. diversifolia (1.62 µm) and the longest in I. clarkei (2.15 µm). In the present inves- tigation, the longest chromosomes were observed in I. ochracea (0.94 to 1.57 µm) whereas the smallest (0.74 to 1.20 µm) in I. marginata f. candida. All the studied spe- cies fell under Stebbins’s karyotype asymmetry class 4A. MCA value was maximum in I. kotschyana (14.65) which indicated greater differences in the centromeric position across the chromosome complement whereas the high- est CVCL in I. corymbosa (14.70 µm) was on account of higher heterogeneity in the length of the chromosome complement. Amongst the Indian Merremia species, cytogeneti- cal data are available for 60% species (after Rice et al., 2015). Lewis et al. (1967) recorded 2n = 28 chromo- somes for M. aegyptia (L.) Urb. whereas 2n = 30 chro- mosomes were reported by Jones (1968) and Pitrez et al. (2008). Vij et al. (1977) observed 2n = 30 chromosomes in M. dissecta (Jacq.) Hallier f. and M. aegyptia. Meio- sis revealed n = 15 bivalents for both M. dissecta and M. aegyptia. Aberrant meiosis was reported in M. aegyptia. The presence of 2n = 28 and 30 chromosomes need to be further investigated. Secondly, this could also be possi- ble due to the existence of two cytotypes in M. aegyptia. Ramanpreet and Gupta (2018) recorded n = 7 bivalents in M. umbellata (L.) Hallier f. Sampathkumar (1979) reported 2n = 32 and 2n =30 chromosomes in M. dis- secta and M. hederacea (Burm.f.) Hallier f., respectively with median and submedian centromeres. R value was 2.5 and 3.3 and the chromosome length ranged from 1.2 µm to 3.0 µm and 1.0 µm to 3.3 µm for M. dissecta and M. hederacea, respectively. We observed 2n = 30 chro- mosomes in M. rhyncorhiza. Accordingly, the R value, i.e. 1.61 was smaller than M. dissecta and M. hederacea. The chromosomes were smaller in size (0.85 µm to 1.38 µm) and had median region centromeres and symmet- rical karyotype. Pitrez et al. (2008) also reported sym- metrical karyotype in M. aegyptia but the chromosomes were with metacentric and submetacentric region cen- tromere with terminal secondary constriction on one of submetacentric pairs. Sampathkumar (1979) also found satellite chromosomes and secondary constrictions in both species M. dissecta and M. hederacea. We could not observe any satellite chromosomes and secondary con- strictions in M. rhyncorhiza. Recently, Ramanpreet and Gupta (2018) carried out meiotic studies on 19 species of Convolvulaceae from Indian hot desert Rajasthan. They studied nine Ipomoea species and reported normal meiosis and high pollen fertility in all studied Ipomoea species. For I. cordatotriloba Dennst., I. triloba and I. sagittifolia Burm.f. a meiotic count of n = 15 bivalents was record- ed for the first time (Ramanpreet and Gupta 2018). They found n = 15 bivalents for eight Ipomoea species and n = 14 bivalents for I. pes-tigridis. This study also confirmed earlier reports on chromosome numbers of Ipomoea. In present investigation, meiotic counts of n = 15 in I. horsfalliae, I. laxiflora, I. marginata f. candida and I. salsettensis and n = 30 in I. ochracea were made for the first time. Meiotic course was normal. Lekhak et al. (2018) also found normal meiosis in I. clarkei and I. diversifolia. Vij et al. (1977) also studied meiosis in Ipomoea and some allied genera. They studied meiosis in 22 Ipomoea species and found normal bivalent for- mation. Most of the Ipomoea species exhibited n = 15 bivalents. I. coccinea L. and I. batatas were reported to have the counts of n = 14 and n = 45, respectively. Irregular anaphases were also observed some species. One B-chromosome was observed in I. mutabilis and I. palmata (white flowered type). We did not find B-chro- mosomes in the meiotic phases. Chromosome data are now lacking only in fourteen Ipomoea and six Mer- remia species in India (Table 3). 64 R.N. Chougule et al. CONCLUSION Comprehensive data on cytogenetics of Ipomoea and Merremia are important to understand the chromo- somal evolution and harness the economic potential. As molecular phylogeny for the Indian taxa is not available, karyological data, particularly chromosome number and information from f luorescent banding or f luorescent in-situ hybridization can help to reveal species interre- lationships. Based on the data of chromosome number for Ipomoea and Merremia presented here and previ- ous reports it is clear that there are two basic chromo- some numbers, i.e. x = 14 and x = 15. Nevertheless, more information on the hitherto studied taxa and confirma- tion of chromosome number in taxa where the count does not confirm to these numbers is needed to under- stand mechanisms underlying evolution in these genera. 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Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Volume 76, Issue 2 - 2023 Firenze University Press Molecular classification of Barbeyaceae (Barbeya oleoides Schweinf.) using four different DNA barcodes Fatima Omari Alzahrani, Sami Asir Al-Robai Mitotic metaphase karyotype of the mosquito Anopheles arabiensis Patton (Diptera: Culicidae) from Kassala State, eastern Sudan Asma Mahmoud Hamza1,*, Sumaya Hussein Elboshra2 Karyotypic analysis of Crucian carp, Carassius carassius (Linnaeus, 1758) from cold waters of Kashmir Himalayas Gousia Jan1, Asim Iqbal Bazaz2, Azra Shah1, Saima Andleeb1, Irfan Ahmad1,*, Durdana Qazi1, Oyas Asimi3, Bilal A. 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