Chromosome Botany (2008) 3: 41-47 © Copyright 2008 by the International Society of Chromosome Botany

Karyomorphological studies of some of Chrysanthemum sensu lato in Egypt

Magdy Hussein Abd El-Twab1, Ahmed Mohammed Mekawy and Mohammad Saad El-Katatny

Department of Botany, Faculty of Science, Minia University, El-Minia City 61519, Egypt

1Author for correspondence: ([email protected]) Received June 8, 2008; accepted September 21, 2008

ABSTRACT. Chromosome numbers and karyotypes are presented in eight taxa in Chrysanhemum senu lato (). Achillea santolina, Cotula barbata and Co. cinerea had new counts for the first time. The chromosome complement of arborescens (2n=18) consists of 14 median and four sub- median chromosomes. Achillea fragrantissima (2n=18) consists of 14 median and four sub- median chromosomes; Ach. santolina (2n=36) consists of 26 median- and ten submedian-centromeric chromosomes. Glebionis coronaria (2n=18) consists of 18 median-centromeric chromosomes; while G. coronaria var. discolor (2n= 18) has 16 median- and two submedian-centromeric chromosomes. Cotula barbata (2n=16) consists of 12 median- and four sub-median-centromeric chromosomes, while Co. cinerea (2n=18) has 14 median- and four sub-median-centromeric chromosomes. The chromosome complement of Matricaria recutita (2n=18) consists of 14 median-, two submedian- and two subterminal-centromeric chromosomes.The longest chromosome complement was Artemisia arborescens while the shortest was Cotula barbata. The length of the tetraploid chromosome complement of Ach. santolina was approximately one third shoter than the diploid Artemisia arborescens, and equal to G. coronaria. KEYWORDS: Achillea, Artemisia, Chromosome, Chrysanthemum, Cotula, Glebionis, Karyotypes, Matricaria

The Asteraceae is an immense family that contains subtribe Anthemidinae represents all the Anthemidae many major ornamental and medicinal (Tang et al. with a paleate receptacle, it is essentially restricted to the 2000). The tribe is one of the largest tribes large Anthemis. The subtribe Chrysantheminae is a of the family with 1741 species predominantly distributed small group distributed in Eurasia, North Africa and in Eurasia, North and South Africa, with fewer species in Macaroesia and traditionally it comprised all of the North America and Australia (Bremer and Humphries Anthemideae species with an epaleate receptacle (Bremer 1993). It is an important tribe that contains medicinal and Humphries 1993). The Chrysantheminae is considered importances. Many species are weeds of cultivated lands, a possible sister group candidate to Anthemidinae. The disease resistant and have other desirable agronomical subtribe Artemisiinae is distributed widely in Northern traits including Artemisia, Chrysanthemum and Matricaria Hemisphere especially in Central and East Asia (Bremer (Silva 2004). The principal taxonomic problems within and Humphries 1993). the tribe are entirely of relationships between genera and Studies on origin and chromosome constitutions of also circumscription of the genera (Bremer and the native polyploid species of Chrysanthemum sensu Humphries 1993). According to Tackholm (1974) lato may contribute to satisfactory phylogenetic and Anthemideae is represented in Egypt by 11 genera and taxonomic treatment of species relationships and the 31 species, several species are very rare and became very constitution and breeding . Chrysanthemum difficult to find them in the natural habitats now a days. sensu lato performs a polyploid series which plays an Chrysanthemum sensu lato consisted of 27 genera has important role in chromosome evolution with the basic been taxonomically placed in the subtribe Chrysantheminae chromosome number of X=9 (Tahara 1915; Nakata and O. Hoffm., the tribe Anthemideae Cass (Shih and Fu Tanaka 1987; Abd El-Twab and Kondo 2003). In plant 1983), while it has been placed in different subtribes in evolutionary studies and breeding there is often a need to the tribe Anthemideae (Bremer and Humphries 1993). discriminate between the genomes of closely related Accordingly, Artemisia has been placed in the subtribe genera or species and to identify the ancestors of Artemisiinae, Achillea and Anacyclus in the Achilleinae, polyploid species (Ørgaard and Heslop-Harrison 1994; Chrysanthemum in the Chrysantheminae and Cotula and Abd El-Twab and Kondo 2003). Matricaria in the Matricariinae. The main centers of This study aims to contribute towards a better distribution of Chrysanthemum in broad sense are two; understanding of the karymorphological relationships one in the Mediterranean area and the other in China and and evolution of Chrysanthemum and closely related Japan. The Chinese and Japanese species have a wide genera of Chrysanthemum sensu lato distributed in chromosomal range from 2n =18 (diploid) to 2n =90 Egypt. (decaploid). They are considered as ancestors of the Mediterranean species (Dowrick 1952). Traditionaly, the

41 42 ABD EL-TWAB ET AL.

Table 1. Chromosome number, length, chromosome complement lengths and locality of the species studied in Chrysanthemum sensu lato distributed in Egypt.

Chromosome length Total Chromosome complement

Chromosome No. µm Lengths µm Locality Species 2n shortest+SD longest+SD shortest longest Mean+SD Artemisia arborescens 18 4.23+0.45 7.32+0.8 84.2 135.8 104.5+6.8 Minia university Achillea fragrantissima 18 2.68+0.2 4.9+0.3 47 73.25 61.67+4.6 Wadi Hagool Achillea santolina 36 1.66+0.2 2.9+0.7 56.3 91.28 79.3+5.4 Burg El-Arab Glebionis coronaria 18 3.28+0.4 5.44+0.2 64.55 95.95 79.2+4.0 El-Minia city Glebionis coronaria var. 18 2.86+0.3 4.31+0.2 63.65 68.6 65.45+2.7 El-Minia city discolor Cotula barbata 16 1.16+0.04 1.86+0.03 22 27.42 24.24+1.1 Minia university Cotula cinerea 18 1.62+0.1 4.35+0.4 46.54 58.87 52.7+8.5 Cairo- Alex. Road Matricaria recutita 18 1.45+0.2 2.56+0.37 25.3 45.2 34.9+1.6 El-Minia city SD: Standard Deviation

MATERIALS AND METHODS variable species (or mixture of species) with gray-green Plant material The plants used in this experiment (Table to silver leaves. It is native to the various habitats of the 1) were Artemisia arborescens L. Achilliea fragrantisma Mediterranean region, where it occurs as a shrub to one (Forssk.) Sch. Bip, Ach. santolina L., Glebionis meter in height. The wild species of Artemisia coronaria (L.) Trehane L., G. coronaria var. discolor D’ arborescens may have originated in North Africa or the Urv, , Cotula barbata DC. Co. cinerea Del, and Middle East (Tucker et al. 1993) and was reported by Matricaria recutita L. The dried plant materials were Delile in the gardens of Egypt (Pickering 1854). preserved in the Herbarium of Botany Department, Artemisia arborescens has been placed in the subtribe Faculty of Science. Minia University, El-Minia City, Artemisiinae (Bremer and Humphries 1993). The present Egypt. count chromosome number of Artemisia arborescens of 2n= 18 (Fig. 1 and 2 A) was convenient with previously Chromosome preparations The method followed Kondo reported counts (Pavone et al. 1981; Ouyahya and Viano et al. (1992) with little modifications as follow: Seeds 1988; Vallés and Torrell 1995). The chromosome were collected and germinated in Petri-dishes. The complement of A. arborences has approximate length of growing root tips were cut (1: 1.5 cm long) and pretreated 104.5 µm (Table 1) and consists of 14 median- and four with an aqueous solution of 0.002M 8-hydroxyquinoline sub-median- centromeric chromosomes. The sat- for 4 h at 4˚C. Fixation was done in acetic-alcohol (1part chromosomes were one pair and not clear in all plates. of glacial acetic acid + 3 parts of 95% ethanol) for 24 h Achillea was established by Linnaeus (1753) and at 4˚C. Maceration was performed in HCl-acetic acid (2 placed by Lessing (1832) and Hoffmann (1894) in sub- parts of 1N HCl + 1part of 45% acetic acid) for 1 min at tribe Anthemideinae, while it was replaced in the sub- 60˚C. The roots were then washed by distilled water for tribe Achilleinae by Bremer and Humphries (1993). In 10 min at room temperature. After that, the meristematic Egypt the genus is represented by two species A. region of the root tip was squashed in a drop of aceto- fragrantissima and A. santolina (Tackholm 1974). In the carmine stain on a slide glass, applied a cover glass on it, present count the chromosome number of Achillea trapped lightly and squashed strongly. Examination of fragrantissima is 2n= 18 (Fig. 1 and 2 B) that is slides was done by Zeiss microscope and photographs for convenient with previously reported count (Badr et al good plates were taken with a digital camera “Ulead 1997), while Achillea santolina is 2n= 36 (Fig. 1 and 2 explorer.” Analysis of chromosome complements and C), which considered as a new count and different from accumulation of data were based on the microphotography. that was reported as 2n=54 (Androshchuk and Kostinenko 1981) with x=9 (Ghaffari 1989). In the RESULTS AND DISCUSSION present study the chromosome complement of Achillea Chromosome numbers and karyotypes of 8 taxa in fragrantissima has approximate length of 61.7 µm, and Chrysanhemum senu lato are presented in Table 1. consists of 14 median and four sub-median-centromeric Achillea santolina, Cotula barbata and Cotula cinerea chromosomes, while Ach. santolina has approximate had new counts for the first time. Six species were length of 79.3µm and consists of 26 median- and ten sub- diploids with 2n= 18, one species was anueopliod with median-centromeric chromosomes. The sat-chromosomes 2n= 16, and one species was tetraploid with 2n= 36. were one pair and two pairs in the chromosome Artemesia arborescens is a morphologically highly complements of Ach. fragrantissima and Ach. santolina, KARYOMORPHOLOGY OF EGIPTIAN CHRYSANTHEMUM 43

Fig.1. Metaphase chromosome plates stained by aceto-carmen. A. Artemisia arborescens. B. Achillea fragrantissima. C. Ach. santolina. D. Glebionis coronaria (yellow legules). E. G. coronaria var. discolor (white-yellow legules). F. Cotula barbata. G. Cotula cinerea. H. Matricaria recutita. Bar= 5 µm for all chromosome figures. 44 ABD EL-TWAB ET AL.

Fig.2. Karyotypes of metaphase chromosomes (A-H). A. Artemisia arborescens. B. Achillea fragrantissima. C. Ach. santolina. D. Glebionis coronaria (yellow legules). E. G. coronaria var. discolor (white-yellow legules). F. Cotula barbata. G. Cotula cinerea. H. Matricaria recutita. Bar= 5 µm. respectively. The chromosome complement of G. coronaria has In Egypt, the genus Glebionis is represented by one approximate length of 79.2 µm and consists of 18 species Glebionis coronaria (yellow ligules) and one median-centromeric chromosomes, while C. coronarium variety G. coronaria var. discolor (yellow-while ligules; var. discolor has approximate length of 65.45 µm and Tackholm 1974). The present study is the first to consists of 16 median- and two sub- median-centromeric represent the count and karyotype of this species and its chromosomes. The sat-chromosomes were one or two variety. The present count of the 2n= 18 (Fig. 1 and 2 D) pairs in the metaphase chromosomes of G. coronaria and is convenient with previously reported count of G. its variety. coronaria (Bhattacharyya 1977; Pavone et al. 1981; The present count of the 2n= 16 of Cotula barbata Razaq et al. 1988; Nirmala and Rao 1986; Vogt and DC. (Fig. 1 and 2 F) had the chromosome complement Oberprieler 1993; Ruíz de Clavijo 1993; Oberprieler and length of 24.3 µm, which consists of 12 median- and four Vogt 1993; Kaul and Kaur 1995; Strother and Watson submedian-centromeric chromosomes, while the present 1997; Carr et al. 1999; Vogt and Aparicio 1999), and of count of 2n=18 of C. cinerea Del, (Figs. 1 and 2G) has G. coronaria var. discolor (Vogt and Aparicio 1999). the chromosome complement length of 52.7 µm, which KARYOMORPHOLOGY OF EGIPTIAN CHRYSANTHEMUM 45 divided into 14 median- and four submedian-centromeric chromosomes of the complement are median-centromeric chromosomes. The sat-chromosomes were one or two chromosomes) and the chromosome complement of pairs among the metaphase chromosomes. Matricaria recutita the most advanced karyotype (14 Matricaria was established by Linnaeus (1753) and median-, two submedian- and two subterminal- placed by Lessing (1832) and Hoffmann (1894) within centromeric chromosomes). The evolution of their extended sub-tribe Chrysanthemineae. Reitbrecht chromosome shape, size, composition, number and (1974) placed it in Matricaria-group with Anthemis and redundancy might result in a wide diversity of Tripleurospermum. Matricaria is represented in Egypt karyotypes. Several models are describing the by three species (Tackholm 1974), this genus already karyotypes evolution (the stasipatric model: White 1973; segregated into several genera of which Chlamydophora the canalization model: Bickham and Baker 1979; the Ehrenb. ex Lessing is represented in Egypt by peripatric speciation model by centric fusion: Baker and Chlamydophora tridentata. The present count of 2n=18 Bickham 1986). of Matricaria recutita L. (Fig. 1 and 2H) was convenient Addressing chromosome evolution by combination of with previously reported count (Matricaria chamomilla: different experimental approaches might reveal new eg; Arora and Madhusoodana 1981; Madhusoodanan and insights, such as the re-interpretation of chromosome Arora 1979, 1984. Matricaria recutita: eg, Rostovtseva number, size and morphology among the species of 1979; Magulaev 1979; Kuzmanov et al. 1986; Peneva et Chrysanhemum senu lato (Kondo and Abd El-Twab al. 1988; Vogt and Aparicio 1999). Matricaria recutita 2002; Abd El-Twab and Kondo 2003). Hence, this has the chromosome complement length of 34.9 µm, study aims to contribute towards a better understanding which divided into 14 median-, two submedian- and two of the cytological evolution of Chrysanthemum and subterminal-centromeric chromosomes. The sat- closely related genera in Chrysanthemum sensu lato chromosomes were one pair among the metaphase distributed in Egypt. We report chromosome numbers for chromosomes. eight taxa (several taxa from more than one accession An especial striking property of satellite (sat) DNA is were tested), three of these chromosome complements its apparent liability in evolution, both in a qualitative were counted for the first time (in Egypt), plus and quantitative sense (Lohe and Roberts 1988). Since, karyotypes for representatives of most of the major a prediction of chromosomal locations of satellites would lineages. The longest chromosome complement was be that closely related satellite sequences show similar Artemisia arborescens while the shortest was Cotula chromosomal distributions. Closely related satellite barbata. The length of the tetraploid chromosome repeats occupy adjacent regions of the chromosomal complement of Ach. santolina was approximately one DNA. Such topological constraints might arise from third shorter than the diploid chromosome complement either functional restrictions or from mode of origin, one of Artemisia arborescens, while was approximately equal from the other (Lohe and Roberts 1988). The taxa in this to the chromosome complement of G. coronaria. The study showed variations in visible numbers of satellites present karyological data are synthesized and used to in each chromosome complements at mitotic metaphase formulate hypotheses concerning the evolution of basic by Carmen staining, probably due to either too much chromosome numbers and the karyotypes within condensation of satellite by 8-hydroxyquinoline and/or Chrysanthemum and related genera. The information buried satellite in chromosome arm or too much that has been presented her is a step towards understanding physically stretched secondary constriction (Kondo et al. the variation in chromosome complements, numbers and 1996). morphlogy. We still in need for more chromosome Karyotype evolutions are one of the most important studies employing both classical and molecular aspects of the whole evolutionary processes (Imai et al. cytogenetic methods, which might be an important 1986) and considered as an isolating mechanism in source of information for analyzing taxonomical speciation and have their own evolutionary trends relationships and evolution of the taxa. Therefore more independent of genetic evolution (Imai et al. 2001). studies are planned on the chromosome complements of Therefore, karyotype evolution generally tends towards the species to clarify and justify the species taxonomical an increasing number and terminal-centromeric relationships, evolution and polyploidization mechanism chromosomes (acrocentric). The opposite tendency, the in Chrysanthemum sensu lato. reduction of chromosome number and formation of median centromeric chromosomes (metacentric) are ACKNOWLEDGEMENT. We thank Prof. Dr. Emeritus M.A. primitive (Imai et al. 2001). Accordingly, the Elnaghy for valuable suggestions and support during the course of this study. karyotype of the tetraploid chromosome complement of Achillea santolina is considered the most advanced LITERATURE CITED karyotype among the presented species. While, among Abd El-Twab, M. H. and Kondo, K. 2003. Physical mapping the diploid species, the chromosome complement of G. of 45S rDNA loci by fluorescent in situ hybridization and coronaria is considered the most primitive karyotype (all Evolution among polyploid Dendranthema species. 46 ABD EL-TWAB ET AL.

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