(Matthiola Incana L.) Flower
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FOLIA HORTICULTURAE Folia Hort. 29/1 (2017): 51-61 Published by the Polish Society DOI: 10.1515/fhort-2017-0006 for Horticultural Science since 1989 ORIGINAL ARTICLE Open access http://www.foliahort.ogr.ur.krakow.pl Meiotic behaviour and morpho-phenological variation in cut stock (Matthiola incana L.) flower Sepideh Famil Irani, Mostafa Arab* Department of Horticulture College of Aburaihan, University of Tehran 20th Km Imam Reza Road, Tehran, Iran ABSTRACT Morpho-phenological and meiotic studies were performed in twelve cultivars of Matthiola incana. All of the cultivars were diploid (2n = 2x = 14) with basic chromosome number x = 7. A number of aneuploid PMCs (n + 1) were observed in plants of two cultivars, named ‘Nobel’ (NB) and ‘Goddess’ (GD), at the diakinesis stage. Trisomic individuals with the frequency of 20% and 5% and (2n + 1 = 15) somatic chromosomes were observed in seeds obtained from single-flowered plants of the cultivars NB and GD, respectively. An additional chromosome was mostly observed in the form of a chain trivalent or a rod univalent. Various meiotic abnormalities were found in all the cultivars to different degrees. In these cultivars, the percentage of cells with meiotic abnormalities was higher in anaphase I. Cytomixis was observed for the first time in Matthiola incana. ANOVA tests revealed significant differences in morpho-phenological characteristics. ‘Nobel’ differs from the others in all of the vegetative features investigated in this study. All the cultivars studied except ‘Nobel’ and ‘Pacific Crimson’ possessed high pollen fertility (> 90%). Five groups of the cultivars based on morpho-phenological features disagree with the clustering of cultivars based on meiotic traits. It is thought that the various morpho-phenological features observed among the cultivars could be due to their different genetic background and not only to meiotic anomalies. Key words: aneuploidy, chiasma frequency, chromosome pairing, pollen fertility, vegetative and floral traits INTRODUCTION plants, it has been observed that mutations in the AGAMOUS (AG) gene cause deactivation of the The stock flower Matthiola( incana L.) is a species gene. So, in an AGAMOUS mutant there are petals of the Brassicaceae family. It is regarded as an instead of the stamen and carpel (Meyerowitz et al. important ornamental plant due to its pleasant 1989). Production, however, is problematic because aroma and beautiful form (Sanchez et al. 2005). of the genetic disorder in the double-flowered plant, This plant exists in two forms: single-flowered and which lacks the stamen and carpel. The double- double-flowered. The double-flowered stock plant is flowered form is sterile and can only be produced popular on the market and that makes it an important from seeds obtained from single-flowered plants export commodity worldwide (Winge 1931, Eid (Johnson 1953, Mizukami and Ma 1997). The et al. 2009). Seeds of the stock flower contain oil seeds of the single-flowered plant produce various rich in linolenic acid (55-65%) that is of medicinal percentages of double-flowered plants, depending importance (Yaniv et al. 1999). In double-flowered on the cultivar. According to reports there are *Corresponding author. Tel.: +98 36041089; e-mail: [email protected] (M. Arab). 52 Meiotic behaviour and morpho-phenological variation in cut stock three types of single-flowered stock plant used as and cytological properties. Double-flowered plants a production base for double-flowered plants. These are sterile, so their meiotic properties cannot be are the “Single” type that, when self-pollinated, studied and no relationship has been determined do not produce double-flowered plants; the between cytogenetic and morpho-phenological second type (“High double”) can produce 25% properties in double-flowered plants. Therefore, in double-flowered plants, and the “Eversporting” this study, double-flowered plants were excluded type produces 40-50% double-flowered plants from the beginning of the experiment and were (Emsweller et al. 1937). Some reports suggest counted only for the purpose of investigating the that the effect of chromosomal abnormality could double-flowered plant rate of each cultivar. be connected with the appearance of double- flowered plants, as well as other morphological MATERIAL AND METHODS properties (Frost and Lesley 1954). Most studies on Plant material chromosomes related to this plant date back 50 years. Based on these studies, the basic chromosomal The meiotic and morpho-phenological study was number (n = 7, 2n = 2x = 14) was reported for performed in twelve cultivars of M. incana listed this plant (Allen 1924, Frost and Lesley 1927). in Table 1. One hundred seeds of each cultivar were Frost (1931) identified aneuploidy in a plant called sown in seedling trays, and at the four-leaf stage, “Snowflake” with 86% double-flowered progenies: the plants were transferred to small plastic pots, cytological investigation determined additional then elimination their requirement for chilling, the chromosomes in this type. Philp and Huskins (1931) seedlings were transferred to 20-cm-diameter pots reported that this additional chromosome had for better development and put in the greenhouse a V shape that was eliminated during meiotic under the same conditions. The average daily division but remained in mitosis. Frost and Lesley temperature in the greenhouse was 15-18ºC, and (1959) reported that an extra chromosome occurred the minimum temperature was 10ºC. All the plants as a consequence of chromosome breakage at of each cultivar were labelled and after recording the first meiotic anaphase. This presented the their morpho-phenological properties and the possibility of more than one additional fragment time of full opening of the inflorescence (flower), of chromosome in the process (Armstrong and collecting buds of the plants was conducted. Huskins 1934). Chromosomal research plays an Analysis of morpho-phenological characteristics important role in studies relating to systematics and In general, 14 morpho-phenological traits related to determinations of genetic diversity (Ortega et al. vegetative and reproductive growth of each plant 1977). There is high diversity among stock flower were measured. This experiment was performed cultivars in terms of morphological properties, such in a completely randomized design (CRD) with that the appearance of double-flowered progenies 20 replicates (single-flowered plants) per cultivar. is variable (Tatsuzawa et al. 2012). However, no systematic study has been done to date to determine Separation of single- from double-flowered the chromosomal characteristics of stock cultivars. plants In this study, we investigated: i) cytological According to Ecker (1993), seedlings with pale traits such as chiasma frequency and chromosome green cotyledons were considered double-flowered pairing, ii) morphological (vegetative and plants. There are two types of cultivars in this reproductive) traits, iii) phenological characteristics study: of three types of stock cultivars: “Eversporting” i). Non-selectable cultivars. In cultivars with no cultivars, named ‘Cinderella Red’ (CR), ‘Goddess selectivity potential (the cultivars NB and ES), White’ (GD), ‘Pacific Crimson’ (PC), ‘Rose Pink’ the separation of single-flowered plants (Fig. 1e) (RP), ‘Lavender Lilac’ (LL), ‘Miracle Mid Blue’ from double-flowered plants (Fig. 1f) was not (MB), ‘Avalanche White’ (AV), ‘Hot Cakes Purple’ possible until the emergence of inflorescence. (HP), ‘White Stock’ (WS) and ‘Deep Blue Stock’ ii). Selectable cultivars. They (all the cultivars, (DB), a “High double” cultivar named ‘Nobel except NB, ES) have the potential to be White’ (NB) with 80% production of double- distinguished (double-flowered from single- flowered plants, and the ”Single” type cultivar flowered plants) at the four-leaf stage by the ‘Esfahan Purple’ (ES), which produces single- colour of the cotyledon. So, individuals with flowered plants. Finally, our intention was to a pale green cotyledon (double-flowered plants) categorize cultivars according to morphological were counted and removed. Sepideh Famil Irani, Mostafa Arab 53 Table1. Data showing twelve stock cultivars with locality, type, meiotic chromosome number, chromosome association, pollen fertility, percentage of double-flowered plants No. of Chromosome associations PFd PDe Cultivar Code Sourcea diakinesis nb Type of stock at diakinesisc (%) (%) cells Cinderella Red CR JPN 378 7 [7II], [5II+1IV], [6II+2I], [5II+4I] 94.7 45 Eversporting [7II], [7II+1I], [5II+1IV], [6II+1III], Nobel White NB JPN 510 7 82.1 80 High double [4II+1III+1IV, [6II+2I], [3II+2IV] [7II], [7II+1I], [6II+2I], [5II+1IV], Goddess White GD NED 491 7 96.4 70 Eversporting [4II+1III+1IV], [6II+1III] Pacific Crimson PC USA 569 7 [7II], [5II+1IV] 89.8 60 Eversporting Rose Pink RP USA 248 7 [7II], [5II+1IV], [3II+2IV] 93.9 50 Eversporting Lavender Lilac LL USA 339 7 [7II], [5II+1IV], [6II+2I] 97.5 45 Eversporting Miracle Mid Blue MB USA 277 7 [7II], [5II+1IV] 95.4 50 Eversporting Avalanche White AV USA 225 7 [7II], [5II+1IV], [3II+2IV] 98.5 60 Eversporting Hot Cakes Purple HP IRN 338 7 [7II], [5II+1IV] 95.3 30 Eversporting White Stock WS IRN 458 7 [7II], [5II+1IV], [3II+2IV] 98.6 40 Eversporting Deep Blue Stock DB IRN 340 7 [7II], [5II+1IV], [3II+2IV] 98.9 35 Eversporting Esfahan Purple ES IRN 297 7 [7II], [5II+1IV] 97.4 00 Single aJPN – Japan, NED – Netherlands, IRN – Iran; bn – haploid number; cchromosome associations were observed at diakinesis (italics – extra chromosome