Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 75(2): 53-58, 2022 Firenze University Press www.fupress.com/caryologia ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-1418 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Citation: Chandra Bhanu Singh, Vijay Kumar Singhal, Manish Kapoor (2022) First record of nucleus migration in premei- otic antherial cells of Saccharum spon- taneum L. (Poaceae). Caryologia 75(2): 53-58. doi: 10.36253/caryologia-1418 Received: September 29, 2021 Accepted: May 20, 2022 Published: September 21, 2022 Copyright: © 2022 Chandra Bhanu Singh, Vijay Kumar Singhal, Manish Kapoor. This is an open access, peer-reviewed article published by Firenze University Press (http://www.fupress.com/caryo- logia) and distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided 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. ORCID CBS: 0000-0001-8337-0629 VKS: 0000-0002-7109-7685 MK: 0000-0002-8349-8910 First record of nucleus migration in premeiotic antherial cells of Saccharum spontaneum L. (Poaceae) Chandra Bhanu Singh1, Vijay Kumar Singhal2, Manish Kapoor2,* 1 University Department of Botany, Tilka Manjhi Bhagalpur University, Bhagalpur – 812 007, Bihar, India 2 Department of Botany, Punjabi University, Patiala-147 002, Punjab, India *Corresponding author. E.mail: jdmanishkapoor@yahoo.com, jdmanishkapoor@pbi. ac.in Abstract. The occurrence of nucleus migration is reported for the first time in a clone (2n = 64) of ‘Thatch’ grass (Saccharum spontaneum L.) of the family Poaceae. Usually, its premeiotic antherial cells are thin walled, uninucleate and without any trace of chro- mosome individuality. However, the cells of those anthers that had been affected from flood water stress conditions were anucleated to hexanucleated in varying frequencies. Out of 2567 cells analyzed, two and three cells were noticed to be connected to each other through a well-defined cytoplasmic channel. The nuclei were observed at various stages of their migration in interconnected cells. The remaining cells exhibited a mosaic of anucleate to hexanucleate cells in varying frequencies with a dominance of binucle- ated condition (43.75%). The anucleate ‘ghost’ cells were much smaller in size than the uninucleate, binucleate and multinucleate cells showing insignificant variation among themselves. The anucleate, binucleate and multinucleate cells appeared to be resulted due to nucleus migration through cytoplasmic channels between two cells. The pres- ence of a nucleus in donor cell united with recipient cell having four nuclei of differ- ent sizes, diminutive anucleate cell in the neighbourhood of uninucleate/trinucleate cell or connected with cytoplasmic channel/pentanucleate cell, and disorganizing cytoplas- mic channel attached with binucleate/ tetranucleate cell witnessed the accomplishment of nucleus migration. This rare phenomenon of nucleus migration seemed to be trig- gered by flood water induced stress and facilitated by feeble cell wall. The variation in sizes of nuclei in multinucleate cells might be due to the transfer of nucleus/nuclei of different size(s). The prominent features of nucleus migration distinguishing it from the cytomixis have been discussed in detail. The syncytes resulted due to nucleus migration might have generated the pollen grains with different genetic constitution resulting into the origin of new intraspecific aneuploids/ polyploids for better adaptability. Keywords: Cytomixis, Nucleus migration, Premeiotic cells, Saccharum spontaneum. INTRODUCTION Flooding stress has been considered as the strong driver of adaptive evo- lution (Jackson and Colmer 2005). In the disturbed habitat of diara land, it 54 Chandra Bhanu Singh, Vijay Kumar Singhal, Manish Kapoor seems to act as the main stress that can trigger syncyte formation leading to the production of new polyploids/ aneuploids. Cytomixis (inclusive of nuclear migration) that operates as the most common pathway of syncyte formation is an important process of evolutionary sig- nificance (Kravets 2012, Mandal et al. 2013, Mursali- mov et al. 2013). The phenomenon is considered to be an efficient mechanism for the production of unreduced (2n) pollen grains and thereby the origin of intraspe- cific polypoids in several plant species (Falistocco et al. 1995, Ghaffari 2006, Kim et al. 2009, Sheidai et al. 2009, Fadaie et al. 2010, Singhal et al. 2010, 2011, 2016, Kaur and Singhal 2012). The polyploids with more adaptabil- ity often thrive better in harsh and disturbed environ- ments (Ramsey and Schemske 1998, Otto and Whitton 2000, Madlung 2013, De Storme and Mason 2014, de Peer et al. 2021) The occurrence of nucleus migration has been observed in the premeiotic cells of anthers of a clone of Saccharum spontanuem L. (English - Thatch grass, Vernacular - Kans, Family - Poaceae) growing under flood water stress conditions. This clone (2n=64) grows profusely in Bhagalpur (Bihar) diara land of Gan- ga basin, where flood is almost a perpetual annual fea- ture (Singh et al. 2018). This is the favourite fodder of buffaloes. Besides, the species is very useful in thatch- ing of roofs and in making various items such as cord- age, ropes, mats, baskets, brooms, etc. (Singh 1997). The Indian sub-continent including Bihar is consid- ered as the centre of greatest evolutionary activity of S. spontanuem L. (Panje and Babu 1960). The occurrence of nucleus migration in the premeiotic cells is the first record for this species, the most valuable germplasm in sugarcane breeding. Fifty-two years ago, chromosome and nucleus migrations were observed during micro- sporogenesis of some radiation-induced mutants of Pisum sativum L. (Gottschalk 1970). The present com- munication aims to clear, enrich and elaborate the con- cept of nucleus migration. MATERIALS AND METHODS The spikelets of a clone of Saccharum spontaneum L. (Voucher specimen number 2180 of Herbarium of Uni- versity Department of Botany, Tilka Manjhi Bhagalpur University, Bhagalpur) growing profusely in Bhagalpur diara land constituted the experimental material of the present investigation. These were collected every alter- nate day from both normal (flood free) and submerged conditions. The collected spikelets from these environ- ments were fixed separately in Carnoy’s fluid (6 etha- nol: 3 chloroform: 1 glacial acetic acid). The fixative was changed daily for 15 days to clear the cytoplasm. Subse- quently, the material was preserved in 70% ethanol and refrigerated at 0ºC until use. Anthers were squashed in 2% acetocarmine and preparations were studied to record abnormalities in premeiotic cells. Photomicro- graphs of unusual premeiotic cells were taken from the temporary preparations. The diameters of visible nuclei were determined from their enlarged photographs (at constant magnification) and the volumes of nuclei were calculated using the formula 4/3 for sphere; where r is the radius (i.e., half of diameter). RESULTS The premeiotic cells from young anthers of a diara clone of ‘Thatch’ grass (Saccharum spontaneum L.) had thin wall and a distinct nucleus. These cells were con- stantly uninucleate in the anthers of normal plants. However, wide variations in number of nuclei were observed in the cells from the anthers of flood affected plants. Out of a total 2567 cells analyzed, two and three cells were found to be connected to each other through a conspicuous cytoplasmic channel respectively, in nine- teen (Figure 1) and two (Figure 2) instances. The nuclei were seen at various stages of their migration in the united cells (Figures 2-3). The remaining cells exhibited a mosaic of anucleate to hexanucleate cells (Figures 4-8) in varying frequencies (Table1) with dominance of binu- cleate condition (43.75%). Among the multinucleate cells, the trinucleate ones were the most frequent, while the pentanucleate were the least common. With respect to size, the anucleate ‘ghost’ cells were much smaller than the nucleate (uninucleate, binucleate and multinucleate) cells which did not show significant variation among themselves. The anucleate, binucleate and multinucleate cells appeared to be the resultant products of nucleus migra- tion, commencing with the development of a well- defined cytoplasmic channel between two cells (Figures 1-6). The possession of one nucleus in the donor cell, which is united with a recipient cell containing four nuclei of different sizes (Figure 3) showed that the cyto- plasmic channel might have paved the way for gradual transference of nucleus/nuclei through it. The presence of a diminutive anucleate donor cell in the neighbor- hood of a trinucleate cell (Figure 1) or uninucleate cell (Figure 2) or connected prominently with a cytoplasmic channel (Figure 2) or pentanucleate cell (Figure 6) and the disorganizing cytoplasmic channel attached with a binucleate cell (Figure 4) or a tetranucleate cell (Figure 5), suggested towards the completion of nucleus migra- 55First record of nucleus migration in premeiotic antherial cells of Saccharum spontaneum L. (Poaceae) tion. The phenomenon of nuclear migration seemed to be induced by abiotic stress due to flood water condi- tions and facilitated by feeble wall character of premei- otic cells resulting into syncyte formation. The shapes of cells were noted to be specific to some extent to their nuclear status as exclusively spherical for anucleate (in free condition) to trinucleate; spherical or ellipsoidal for tetranucleate; spherical or irregular for pentanucleate and solely irregular for hexanucleate con- ditions (Table 1). The uninucelate and binucleate cells displayed the presence of only large sized nucleus and nuclei accordingly in them. Irrespective of their shapes, the multinucleate cells had nuclei of different sizes in them (Table 2). These nuclei were arbitrarily classified as large (above 50 µm3), medium (above 20 µm3 and up to 50 µm3) and small (up to 20 µm3). The presence of equal sized nuclei in binucleate cells and unequal sized nuclei in multinucleate cells seems to have been derived from similar and dissimilar types of cells. This became evi- dent from the existence of an anucleate ghost cell and a binucleate cell that was connected with a trinucleate cell (having one nucleus of each category) through a cyto- plasmic channel for nuclear migration (Figure 1). The size of nucleus contained in uninucleate cells showed approximation to the dimension of large sized nuclei possessed by tetranucleate cells, whose medium sized nuclei were almost of the magnitude of medium sized nuclei of trinucleate cells. Figure 5 indicates the presence of a tetranucleate cell possessing different sized nuclei, which might have resulted due to nucleus migrations from different types of donor cells. DISCUSSION Ever since the first report of syncytes by Gates and Reese (1921), these multinucleate cells have been report- Figures 1-8. Premeiotic antherial cells of Saccharum spontaneum L. showing nucleus migration and its products. 1. Two cells con- nected through a conspicuos cytoplasmic channel (arrowed) in between and a small anucleate ghost cell (arrowhead) present on upper side. 2. Two distinct uninucleate cells linked through a cyto- plasmic channel (arrowed) and a small deformed anucleate ghost cell (arrowhead) united mid way with the channel – discharged nucleus (double arrowed) entering into a cell to make it binucleate; another small spherical free anucleate ghost cell (arrowhead) and a large binucleate cell discernible on lower side. 3. Two cells intercon- nected with each other through a cytoplasmic channel (arrowed): Donor cell possessing one nucleus and recipient cell containing four unequal sized nuclei; a binucleate cell visible above. 4. Uni, bi and trinucleate cells; a binucleate cell united with a disorganizing cyto- plasmic channel (arrowed). 5. Tetranucleate cells showing marked variation in the size of nuclei and one tetranucleate cell showing attachment with a disorganizing channel (arrowed). 6. A tetra- nucleate cell in left and an elongated cum curved anucleate ghost cell (arrowhead) attached with a pentanucleate cell (having uneven sized nuclei) in right. 7. A deformed pentanucleate cell having nuclei of different sizes. 8. Bi and hexanucleate cells: Curved shape of hexanucleate cell remarkable. Bar = 20μm. Table 1. Type, shape, and frequency of flood affected premeiotic antherial cells* of Saccharum spontaneum L. Cell type Shape Number and Frequency (%) Anucleate Spherical 31(01.21) Uninucleate Spherical 234(09.12) Binucleate Spherical 1123(43.75) Trinucleate Spherical 954(37.16) Tetranucleate Spherical/ Ellipsoidal 165(06.43) Pentanucleate Spherical/ Irregular 16(00.62) Hexanucleate Irregular 44(01.71) *Among 2567 cells, two and three cells connected to each other through a conspicuous cytoplasmic channel respectively in nineteen and two cases for nucleus migration. 56 Chandra Bhanu Singh, Vijay Kumar Singhal, Manish Kapoor ed in several plant species by different workers. The syncytes in plants are usually formed through arche- sporial error, cell fusion and nuclear migration. Out of these, nuclear migration occurs through the transfer of whole chromatin material or nucleus from one cell to an adjacent cell. The migration of partial or total chro- mosome, referred to as “Cytomixis”, is a widespread phenomenon and has been reported in a large number of plant species (Kravets 2012, Mandal et al. 2013, Mur- salimov et al. 2013, Rana et al. 2014, 2015, Kumar et al. 2015, Reis et al. 2015, Bhat et al. 2017, Mandal and Nandi 2017, Singhal et al. 2018, Paez et al. 2021). How- ever, nucleus migration has been observed only in pol- len mother cells of radiation-induced ‘pea’ (Gottschalk 1970). The occurrence of nucleus migration through a conspicuous cytoplasmic channel as observed in the present study appears to be closely allied with the well- known phenomenon of cytomixis through cytomictic channel. Both these phenomena seem to be homolo- gous but indeed separate from each other at least in mode of operation. Cytomixis involves chromatin trans- fer between proximate cells (Kamra 1960, Omara 1976, Belluci et al. 2003, Singhal et al. 2009, 2018, Himshikha et al. 2010, Rana et al. 2013, Kumar and Singhal 2016, Kumar et al. 2016, 2017, Bhat et al. 2017, Mandal and Nandi 2017). Owing to this unique feature, it is also called as the phenomenon of intercellular chromatin transmigration (Kumar and Naseem 2013, Kumar and Choudhary 2016, Dwivedi and Kumar 2018, Khan et al. 2018, Kumar and Singh 2020). Akin to cytomixis, nucle- us migration requires the passage of a whole nucleus from donor cell to recipient cell (Gottschalk 1970, Patra et al. 1987). According to Kihara and Lilienfeld (1934), the term “Cytomixis” should be used to designate only the transit of structureless chromatin drops (“Ueber- tritte structurloser chromatintrophen”). Keeping in view the above-mentioned limitation placed on the use of this term, Gottschalk (1970) designated the phenom- enon of moving chromosome and nucleus with normal structure as “chromosome and nucleus migration” rather than cytomixis. The denotation of migration of chromo- somes and nuclei, respectively as nuclear chromosome migration and migration of nucleus by Patra et al. (1987) may be a circumstantial consideration that cytomixis and nucleus migration are apart from each other. There appears no hitch in corroborating the event of shift of structured nucleus as evident in the present case as “nucleus migration”. The uniformity in the size of uninucleate, binucle- ate and multinucleate cells in the presently studied grass suggests that only nuclei pass through cytoplasmic chan- nels. Unlike this, cytomixis involves the transfer of both chromatin/ chromosome(s) along with cytoplasm as well as other cell organelles through cytomictic channels (Risueno et al. 1969, Romanov and Orlova 1971, Mur- salimov and Deineko 2011, Kumar and Choudhary 2016, Kumar and Singh 2020). This becomes apparent from the increase in the size of cytomictic products/ recipient cells, as has also been recorded by Sarbhoy (1980) and Singh et al. (1989, 1990). Thus, nucleus migration has been treated here as a separate process different from cytomixis. The possible causes of syncyte formation in plants include effect of chemicals, X-rays, temperature, mois- ture stress, viral infection, culture conditions or genetic factors. In the presently studied grass the syncyte for- mation could be attributed to flood water induced stress conditions prevailing in the diara land. The products of Table 2. Data on number of nucleus or nuclei per cell and dimension of different sized nuclei in flood affected premeiotic antherial cells of Saccharum spontaneum L. Nucleus or nuclei/ Cell Different sized nuclei / Cell Diameter of nucleus Mean ± S.D. (µm) Volume of nucleus (µm3) 1 Large - 1 6.01 ± 0.20 118.89 2 Large - 2 5.04 ± 0.68 67.06 3 Large - 1 5.68 ± 0.70 95.99 Medium - 1 3.47 ± 0.18 21.89 Small - 1 2.33 ± 0.37 06.63 4 Large - 2 6.25 ± 0.28 127.88 Medium - 2 3.48 ±0.29 23.43 5 Large - 3 4.68 ± 0.45 53.69 Small - 2 2.17 ± 0.21 05.35 6 Medium - 3 3.80 ± 0.39 28.74 Small - 3 2.26 ± 0.33 06.05 57First record of nucleus migration in premeiotic antherial cells of Saccharum spontaneum L. (Poaceae) such syncytes might have produced the pollen grains with different genetic constitution, which through chance fusion might have resulted in the production of polyploids/ aneuploids for better adaptability. 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Syncytes during male meiosis resulting into 2n pollen grain formation in Lindelofia longiflora var. falconeri. J Syst Evol. 49: 406-410. Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Volume 75, Issue 2 - 2022 Firenze University Press Cytogenetic Studies of Six Species in Family Araceae from Thailand Piyaporn Saensouk1, Surapon Saensouk2,*, Rattanavalee Senavongse2 Effect of Ag Nanoparticles on Morphological and Physio-biochemical Traits of the Medicinal Plant Stevia Rebaudiana Sherzad R. Abdull, Sahar H. Rashid*, Bakhtiar S. Ghafoor, Barzan S. 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Royle leaves extract against nickel nitrate Made Pharmawati1,*, Ni Nyoman Wirasiti1, Luh Putu Wrasiati2 Chromosomal description of three Dixonius (Squamata, Gekkonidae) from Thailand Isara Patawang1, Suphat Prasopsin2, Chatmongkon Suwannapoom3, Alongklod Tanomtong4, Puntivar Keawmad5, Weera Thongnetr6,* First Report on Classical and Molecular Cytogenetics of Doi Inthanon Bent-toed Gecko, Cyrtodactylus inthanon Kunya et al., 2015 (Squamata: Gekkonidae) in Thailand Suphat Prasopsin1, Nawarat Muanglen2, Sukhonthip Ditcharoen3, Chatmongkon Suwannapoom4, Alongklod Tanomtong5, Weera Thongnetr6,* Evaluation of genetic diversity and Gene-Pool of Pistacia khinjuk Stocks Based On Retrotransposon-Based Markers Qin Zhao1,*, Zitong Guo1, Minxing Gao1, Wenbo Wang1, Lingling Dou1, Sahar H. Rashid2 A statistical overview to the chromosome characteristics of some Centaurea L. taxa distributed in the Eastern Anatolia (Turkey) Mikail Açar1,*, Neslihan Taşar2 Cytotoxicity of Sunset Yellow and Brilliant Blue food dyes in a plant test system Elena Bonciu1, Mirela Paraschivu1,*, Nicoleta Anca Șuțan2, Aurel Liviu Olaru1