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Molecular Evidence for an Interfamilial Laminarialean Cross

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Molecular Evidence for an Interfamilial Laminarialean Cross European Journal of Phycology ISSN: 0967-0262 (Print) 1469-4433 (Online) Journal homepage: https://www.tandfonline.com/loi/tejp20 Molecular evidence for an interfamilial laminarialean cross Michael Liptack & Louis Druehl To cite this article: Michael Liptack & Louis Druehl (2000) Molecular evidence for an interfamilial laminarialean cross, European Journal of Phycology, 35:2, 135-142, DOI: 10.1080/09670260010001735721 To link to this article: https://doi.org/10.1080/09670260010001735721 Published online: 03 Jun 2010. Submit your article to this journal Article views: 149 Citing articles: 15 View citing articles Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tejp20 Eur. J. Phycol. (2000), 35: 135–142. Printed in the United Kingdom 135 Molecular evidence for an interfamilial laminarialean cross MICHAEL K.LIPTACK1,2 AND LOUIS D. DRUEHL1,2 "Bamfield Marine Station, Bamfield, B.C., Canada V0R 1B0 #Department of Biological Sciences, Simon Fraser University, Burnaby, B.C., Canada V5A 1S6 (Received 11 June 1998; accepted 7 December 1999) When generating kelp hybrid crosses in vitro, small blades are frequently seen in single-sex cultures of microscopic gametophytes as well as in putative hybrid crosses. We demonstrate a polymerase chain reaction (PCR)-based method that quickly determines the presence\absence of possible parental genomes in the observed blades. ITS1 nrDNA fragments that varied in length in a species-specific way were amplified. Amplification products from possible hybrid blades, apogamic blades and parental gametophytes were compared on agarose gels. Using these methods, we were able to identify a number of apogamic blades (i.e. formed directly from gametophytic cells) as well as a true hybrid sporophyte blade in crosses between Alaria marginata (Alariaceae) and Lessoniopsis littoralis (Lessoniaceae), members of two separate families of the Laminariales. To our knowledge this is the first genotypic confirmation of a hybrid kelp. Key words: Alaria marginata, hybridization, ITS1, Lessoniopsis littoralis, nrDNA, PCR, rDNA Introduction taxa is supported by the observation that all members of the Laminariales studied by Mu$ ller et al. (1985) displayed Hybridization, the crossing of two individuals from sperm chemotaxis based on lamoxirene. different populations, species or genera, has been an Kelps exhibit a complicated life history beginning with important tool in understanding evolution and taxonomy. dioecious microscopic gametophytes, and syngamy In practice, the ability to hybridize is often used as a judge giving rise to macroscopic diploid sporophytes. After a of relatedness (Lewis, 1996b). Versions of the biological period of rapid size increase, the large sporophytes species concept (Dobzhansky, 1937; Mayr, 1963), or meiotically generate motile spores (meiospores) that give various corollaries (Mallet, 1995), often at least partly rise to gametophytes after recuitment. define separate species as those that cannot form viable Doubts concerning the possible crosses arise because a hybrids. Certain groups of organisms, such as some gametophytic or sporophytic morphology does not tracheophytes (higher plants), seem easily to form natural necessarily correspond to a given ploidy level (Nakahara hybrids among closely related taxa, whereas hybridization & Nakamura, 1973; Le Gal et al., 1996). Processes such as is much rarer in animals (Levin, 1979). In the red algae, autodiploidization (a spontaneous doubling of chromo- hybrids of species in Polysiphonia and Mastocarpus (as some number) and apospory (the generation of gameto- Gigartina) have been described (Rueness, 1973; West et al., phytes without the production of spores) can give rise to 1978). However, cellular aspects of hybridization involve diploid gametophytes. Apogamy (the generation of complex processes, with a myriad of causes of lack of sporophytes without the production of gametes) and development, sterility, or death of the hybrid (Lewis, parthenogenesis\androgenesis (development of sporo- 1996b). phytes from unfused gametes) can generate haploid Hybridization has been examined at a number of levels sporophytes and other unusual ploidy levels (for review in the Laminariales. Interspecific hybrids seem quite see Lewis, 1996a). Small blades are frequently observed in common in some genera, such as Laminaria and Macro- cultures of male and female gametophytes of different cystis (Lewis, 1996b). Intergeneric hybrids (e.g. between species. It is generally impossible to determine whether Macrocystis and Pelagophycus) also have been identified in these result from hybridization, parthenogenesis or andro- the field or produced artificially by crossing (Sanbonsuga genesis, apogamy, or some developmentally arrested & Neushul, 1978; Coyer et al., 1992; Lewis & Neushul, partial hybridization. Recently, Le Gal et al. (1996) have 1995). Even hybrids between members of different kelp shown that unisexual gametophyte cultures as well as families have been reported (Tokida et al., 1958; Cosson & sporophytic or gametophytic growth forms can be of Olivari, 1982). The possibility that true hybridization various ploidy levels in the Laminariales. A reliable might occur between members of different kelp higher genotypic test for the nature of the tissues in question is needed to distinguish true from putative hybids. Correspondence to: M. Liptack. e-mail: tenufolia!hotmail.com or The Internal Transcribed Spacer 1 (ITS1) region of the mliptack!sfu.ca nrDNA cistron was chosen for developing such a test. In Published online 03 Jun 2010 M. K. Liptack and L. D. Druehl 136 most eukaryotes, the nrDNA cistron occurs as tandemly Table 1. Clone, species, and general morphology of plants of repeated units of usually several hundred copies approxi- Alaria marginata and Lessoniopsis littoralis from which DNA was mately proportional in number to genome size (Li, 1983). extracted. Sequences for the highly conserved flanking 18S and 5.8S genes in kelps are currently available from GenBank. Species Strain(s) used Morphology Only the ITS1 sequences show variation amongst the recognized kelp species and genera and can be used Clones Alaria marginata Am1,1 female Gametophyte to investigate the various possible crosses within the Alaria marginata Am1,2 female Gametophyte Laminariales (Saunders, 1991). Alaria marginata Am1,3 female Gametophyte We present here a method that at least partially Alaria marginata Am1,1 male Gametophyte addresses the difficulties encountered in assessing true Alaria marginata Am1,2 male Gametophyte Alaria marginata Am1,3 male Gametophyte hybrids by determining whether parental genomes are Lessoniopsis littoralis L1,1 female Gametophyte present in the hybrid offspring. We demonstrate the Lessoniopsis littoralis L1,2 female Gametophyte efficacy of this method on small blades grown in cultures Lessoniopsis littoralis L1,3 female Gametophyte containing attempted hybrid crosses between a member Lessoniopsis littoralis L1,1 male Gametophyte Lessoniopsis littoralis L1,2 male Gametophyte of the Lessoniaceae, Lessoniopsis littoralis, and a member of Lessoniopsis littoralis L1,3 male Gametophyte the Alariaceae, Alaria marginata. Self-crosses Alaria marginata Am1,1 femaleiAm1,1 Sporophyte male Alaria marginata Am1,2 femaleiAm1,2 Sporophyte Materials and methods male Cultures of Alaria marginata Postels & Ruprecht and Alaria marginata Am1,3 femaleiAm1,3 Sporophyte male Lessoniopsis littoralis (Tilden) Reinke used in this study Lessoniopsis littoralis L1,1 femaleiL1,1 male Sporophyte (Table 1) were from our own collection. A culture Lessoniopsis littoralis L1,2 femaleiL1,2 male Sporophyte identification scheme was developed for all taxa used for Lessoniopsis littoralis L1,3 femaleiL1,3 male Sporophyte crossing experiments. Cultures were identified by a code Attempted intergeneric crosses A. marginataiL. littoralis Am1,1 femaleiL1,1 Sporophyte consisting of the taxon (e.g. Am for Alaria marginata) male followed by the number of the sporophyte from which the A. marginataiL. littoralis Am1,2 femaleiL1,2 Sporophyte gametophyte was collected (e.g. 1), separated by a comma male from a numeral to differentiate various gametophytes A. marginataiL. littoralis Am1,3 femaleiL1,3 Sporophyte male from the same sporophyte (e.g. 2), followed by the sex A. marginataiL. littoralis Am1,1 maleiL1,1 Sporophyte of the gametophyte (either male or female). Note that the female actual Lessoniopsis littoralis culture codes in the Bamfield A. marginataiL. littoralis Am1,2 maleiL1,2 Sporophyte Marine Station collection begin with ‘Ll’, as opposed to female A. marginataiL. littoralis Am1,3 maleiL1,3 Sporophyte ‘L’ which is utilized here for clarity. female Both species were initially isolated from Barkley Sound, B.C., Canada on Crosses the 18th of January, 1990 and maintained in culture at the Bamfield Marine Station by L. Druehl. Note that the actual Lessoniopsis littoralis All equipment used in manipulating and storing gameto- cultures in the Bamfield Marine Station collection begin with the two phytes was initially soaked in 500 mM HCl overnight, letter taxa designation ‘Ll’ (e.g. Ll1,1), as opposed to ‘L’ which is utilized rinsed thoroughly in distilled water, and then autoclaved here for readability. # for 30 min at 1n1kgcm− and 121 mC. Three inde- pendently isolated male and female gametophytes of A. marginata and L. littoralis from previously generated individual of the opposite sex or other taxon used in the single-sex, unialgal cultures were ground
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