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Cytogenetic and Phylogenetic Review of the Genus Lachenalia

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Cytogenetic and Phylogenetic Review of the Genus Lachenalia ® Floriculture and Ornamental Biotechnology ©2012 Global Science Books Cytogenetic and Phylogenetic Review of the Genus Lachenalia Riana Kleynhans1,2* • Paula Spies2 • Johan J. Spies2 1 Agricultural Research Council (ARC), Roodeplaat Vegetable and Ornamental Plant Institute (VOPI), Private Bag X293, Pretoria 0001, South Africa 2 Department of Genetics (116), University of the Free State, P.O. Box 339, Bloemfontein 9300, South Africa Corresponding author : * [email protected] ABSTRACT The genus Lachenalia (family Asparagaceae), endemic to southern Africa, is a horticultural diverse genus, with many species featuring in the red data list of southern Africa. The extensive morphological variation within some species complicates species delimitation and has led to taxonomic confusion. The genus is utilised in a breeding programme where cytogenetic and phylogenetic information is important for the development of breeding strategies. Chromosome numbers of 89 species have been recorded in literature, with 2n = 10 to 56 and n = 5 to 28. B-chromosomes have been described in some species. Basic chromosome numbers include x = 5, 6, 7, 8, 9, (probably 10), 11, (probably 12), 13 and (probably 15). Polyploidy was reported in 19 taxa (23%), and is most common in the x = 7 group. Molecular cytogenetic studies using 5S rDNA, 18S rDNA probes and DAPI staining, as well as molecular systematic studies using trnL-F and ITS1- 2 were used to assess the phylogeny of the genus. All these studies indicated that species with the same basic chromosome number are closely related. The one deviation is that it appears as if there are two separate groups within the x = 7 group. The cytogenetic and molecular studies are further supported by breeding studies, where improved results are generally obtained from crosses within a phylogenetic group or between closely related groups. This review of the literature reveals how different studies obtain similar results regarding the phylogenetic relationships within the genus and how these results can be utilized to improve breeding strategies. It also accentuates that further multidisciplinary studies are needed to solve the evolutionary history of the complex genus Lachenalia. _____________________________________________________________________________________________________________ Keywords: chromosome numbers, cladograms, cross-ability, phylogeny, polyploidy Abbreviations: APG, Angiosperm Phylogeny Group; atpB, ATPase beta chain; DAPI, 4',6-diamidino-2-phenylindole; FISH, Fluores- cent in situ hybridization; ITS1-2, Internal transcribed spacer 1 and 2; MEGA, Molecular Evolutionary Genetics Analysis; n, gametic chromosome number; RAPD, Random amplified polymorphic DNA; rbcL ribulose bisophosphate carboxylase (large); SANBI, South African National Biodiversity Institute; trnL, leucyl-transfer RNA intron; trnF, phenylalanine-transfer RNA; VOPI, Vegetable and Orna- mental Plant Institute; x, basic chromosome number; 2n, somatic chromosome number; 5S rDNA and 18S rDNA, 5S and 18S ribosomal DNA CONTENTS INTRODUCTION........................................................................................................................................................................................ 98 CYTOGENETIC STUDIES......................................................................................................................................................................... 99 Chromosome counts ................................................................................................................................................................................ 99 Chromosome morphology ..................................................................................................................................................................... 101 Basic chromosome numbers and polyploidy ......................................................................................................................................... 105 Meiotic studies....................................................................................................................................................................................... 106 PHYLOGENETIC STUDIES .................................................................................................................................................................... 106 The phylogenetic position of Lachenalia .............................................................................................................................................. 106 Phylogeny within the genus................................................................................................................................................................... 107 CROSS-ABILITY IN LACHENALIA ........................................................................................................................................................ 107 COMPARISON BETWEEN CROSS-ABILITY, CYTOGENETIC AND MOLECULAR DATA ............................................................ 108 Basic chromosome numbers and cladograms ........................................................................................................................................ 108 Basic chromosome number and cross-ability ........................................................................................................................................ 108 Evolution and relatedness of different basic chromosome numbers ...................................................................................................... 109 Existence of basic chromosome numbers .............................................................................................................................................. 112 Existence of hybrid species ................................................................................................................................................................... 113 CONCLUSION.......................................................................................................................................................................................... 113 ACKNOWLEDGEMENTS ....................................................................................................................................................................... 114 REFERENCES........................................................................................................................................................................................... 114 _____________________________________________________________________________________________________________ INTRODUCTION includes the former genus Polyxena (Manning et al. 2004). Lachenalia is a horticultural diverse genus, with a distribu- The genus Lachenalia Jacq. f. ex Murray, previously a tion range extending from the south-western coast of Nami- member of the family Hyacinthaceae (Manning et al. 2004; bia, southward throughout the Northern, Western and Eas- Duncan and Edwards 2006, 2007), but since 2009 reclas- tern Cape provinces of South Africa (Duncan 1998). One sified under the family Asparagaceae Juss. (APG III group species extends as far inland as the south western part of the 2009), is endemic to southern Africa. The genus now also Free State Province (Duncan 1996). Of the 126 species and Received: 16 December, 2010. Accepted: 20 December, 2012. Invited Review Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 98-115 ©2012 Global Science Books Fig. 1 Morphological variation in Lachenalia in the greenhouse. subspecies described, 10% are endangered, 17% are vul- and appearance, collectors have recognized the horticultural nerable, 2% are considered to be near threatened, 6% are potential of the genus for centuries (Duncan 1988; Du critically rare, 9% are rare and 2% are declining (SANBI Plessis and Duncan 1989; Kleynhans 2009, 2011; Reinten 2009). et al. 2012). The huge phenotypic variation was also the The genus is geophytic, deciduous and is usually winter most important reason for the initiation of a breeding prog- growing. The centre of diversity is in the Worcester grid ramme at the Agricultural Research Council in South Africa. (3319) in the Western Cape province of South Africa, with This led to the production of various hybrids and the intro- species diversity decreasing toward the eastern and northern duction of new products to the international pot plant mar- parts of its range (Duncan 2005). Although Lachenalia spe- ket (Fig. 3) (Kleynhans 2006). cies like L. bulbifera and L. obscura are widely distributed, The variability of the genus in terms of morphology and a substantial number of species (e.g. L. moniliformis, L. cytogenetics, however, lead to specific challenges for the mathewsii) have a restricted distribution, contributing to the breeding of new cultivars. Both incompatibility and other vulnerability of these species (Duncan 1998). isolation barriers exists (Kleynhans and Hancke 2002). A Lachenalia occurs in a wide range of habitats, ranging large number of inter-species crosses are unsuccessful from arid to high rainfall areas. Lachenalia rubida for (Kleynhans et al. 2009) and future breeding progress is example always grows in deep, pure sand often very close dependent on information about the genetic variation in the to the sea, whilst a species like L. campanulata on the other genus. Results generated from cytogenetic and phylogenetic hand is found in heavy soil at
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