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      Species name: Cardoon(Cynara cardunculus L.)
      Overview
    Cynara cardunculus L. belongs to the Asteraceae family (ex Compositae) and is native to the Mediterranean Basin. It includes three botanical taxa: the globe artichoke (var. scolymus L), the cultivated cardoon (var. altilis DC) and the wild cardoon [var. sylvestris (Lamk) Fiori]. Being a cross-pollinated diploid species (2n = 2x = 34) with proterandrous and asynchronous sexual maturity, C. cardunculus harbours a highly heterozygous genetic background (Basnitzki and Zohary, 1994; Portis et al., 2005a; Scaglione et al., 2009). Wild cardoon is the ancestor of both cultivated C. cardunculus forms (Rottenberg and Zohary, 1996; Pignone and Sonnante, 2004) , which evolved independently under the influence of distinct anthropogenic selection criteria: globe artichoke for its immature inflorescences (capitula or heads), consumed either as fresh or industrial-processed product, while cardoon for its fleshy leaves. The three subspecies remain fully cross-compatible with one another, and their F1 hybrids are fertile.

    Italy has been claimed as the centre of globe artichoke domestication, which occurred about 1st century AD (Mauro et al., 2009; Portis et al., 2005a), and it harbours the richest cultivated primary gene-pool well-fitting local climatic conditions and consumers’ preferences. Being grown on a regional scale (Mauromicale and Ierna, 2000), cultivated globe artichoke germplasm accounts for more than 120 varietal types, which can be classified on the basis of the harvest time (early or late) or capitulum traits, like dimension, shape, presence/absence of spines, pigmentation of the outer bracts (Basnitzki and Zohary, 1994; Lanteri and Portis, 2008), defining four main varietal types: “Spinosi”, “Catanesi”, “Romaneschi” and “Violetti” (see pictures). In recent years some seed (achenes) propagated varieties are being diffusing in cultivation, but vegetative propagation, by means of basal and lateral offshoots (either semi-dormant or actively growing), or stump pieces, has been adopted for centuries, and it is still largely prevalent in most of the varietal types and local landraces.

    Globe artichoke contributes significantly to the Mediterranean agricultural economy, with an annual production of about 750Mt from over 80kha of cultivated land and with an annual turnover exceeding 500M dollars. Italy is the leading world producer, (486,600 tons/year, followed by Spain, Egypt, but; its cultivation is also spread and increasing in South America, North Africa, China and USA (FAOSTAT 2009, http://faostat.fao.org).

    Cultivated cardoon is usually raised from seed and handled as annual crop; its cultivation is much less widespread than that of the globe artichoke and the crop remains of regional importance in Spain, Italy and the south of France, where it is used in traditional dishes. The edible parts of the plant are the fleshy stems, which are typically collected in late autumn-early winter. Before collection, stems are tied together, wrapped in straw and buried for about three weeks in order to accentuate the flavour. A study based on microsatellite and AFLP-profiling of the most widely grown Italian and Spanish local varieties showed that they form two separate gene-pools and that a considerable level of within-varietal type genetic diversity is present (Portis et al., 2005b). Cultivated cardoon has also been promoted as a source of lignocellulosic biomass (Encinar et al., 2002a; Encinar et al., 2002b; Foti et al., 1999; Ierna and Mauromicale, 2010) and its seed oil is suitable for both comestible and bio-fuel end-uses (Curt et al., 2002; Lapuerta et al., 2005).

    Many studies support the important role of globe artichoke in human nutrition, due to its high content of nutraceuticals like inulin and in particular mono-caffeoylquinic (four isomers) and dicaffeoylquinic (six isomers) (Lombardo et al., 2010; Pandino et al., 2010; Pandino et al., 2011; Schutz et al., 2004). Furthermore, in various pharmacological test systems, both globe artichoke and cultivated cardoon extracts have shown hepatoprotective, anticarcinogenic, antioxidative, antibacterial anti HIV, bile–expelling and urinative activities as well as the ability to inhibit cholesterol biosynthesis and LDL oxidation (Lattanzio et al., 2009).

      Resources
    • Microsatellite primer pairs (2311): available online;
    • Primer pairs and full statistics on 300 microsatellite loci, experimentally assayed: available online;
    • C. cardunculus ESTs (36,323 Sanger reads) on GenBank as of 12/2012: available here
    • 454-EST library for C. cardunculus var. scolymus (‘Romanesco C3’ genotype)- 184Mbp – not yet available online;
    • 454-EST library for C. cardunculus var. altilis (‘A41’ genotype) - 246 Mbp – not yet available online;
    • 454-EST library for C. cardunculus var. sylvestris (‘Creta4’genotype) -263 Mbp – not yet available online;
    • C. cardunculus transcripts/unigenes information, based on 454 de novo assembly – available online;
    • Illumina paired-ends cDNA (2 x 76bp) of eight C. cardunculus representative genotypes – 6.2 Gbp – not yet available online .
      Genetic maps
    The first genetic map of globe artichoke were based on a cross between the two globe artichoke genotypes ‘Romanesco C3’ and ‘Spinoso di Palermo’, by applying a two way pseudo-test cross approach (Lanteri et al., 2006; Acquadro et al., 2009;). New maps have lately been generated from a population of F1 progeny involving the cross ‘Romanesco C3’ by the cultivated cardoon genotype ‘Altilis 41’ (Portis et al., 2009). The cultivated cardoon map comprised nearly 200 loci, falling into 17 major linkage groups (LGs) and spanning just over 1,015 cM, while the globe artichoke one featured 326 loci arranged into 20 major LGs, spanning about 1,486 cM. Recently, thanks to the integration of a large number EST-SSR loci developed within the CGP (Scaglione et al., 2009), the first SSR-based consensus map of C. cardunculus (‘Romanesco C3’ x ‘Altilis 41’ cross) was constructed, providing a framework of 17 LGs which anchors 228 sequence-based features (Portis et al. in press). An EST-SSR based consensus map for Cynara cardunculus has been recently submitted for publication.
      Images of Cynara cardunculus
      
      Figure 1: Four-varietal types classification on the base of head traits. A) "Spinosi" type, B) "Catanesi" type, C) "Romaneschi" type and D) "Violetti" type.
      
      Figure 2: Parents of mapping progenies; A) "Romanesco C3" genotype (globe artichoke), B) "Altilis 41" genotype (cultivated cardoon), C) "Creta 4" genotype (wild cardoon).
      
      Figure 3: Phenotypic variability in the F1 progeny obtained by crossing "Romanesco C3" (globe artichoke) with "Altilis 41" (cultivated cardoon).
      
      Figure 4: Phenotypic variability in the F1 progeny obtained by crossing "Romanesco C3" (globe artichoke) with "Creta 4" (wild cardoon).
      CGP Activities
    The CGP has been involved in the generation of genomic resources, marker development and germplasm characterization for Cynara cardunculus. This has included the generation of 36,321 Sanger reads from globe artichoke ("Green Globe" varietal type) which have been exploited in the development of ca. 240 EST-SSR markers. The latter made it possible to construct the first SSR-based consensus map of the species (Portis et al., submitted). Moreover, 454 and Illumina GA cDNA sequencing of 11 genotypes (including three mapping parents) led the de novo assembly of a reference transcriptome of C. cardunculus, together with the generation of a 195k SNPs database.

    Single nucleotide polymorphisms (SNPs), identified as being informative in F1 mapping progenies has been selected to design a 768-plex Goldengate assay. This will allow to improve map saturation and will provide the basis for QTL identification for horticultural and biomass-related traits. Many of the selected SNPs reside in genes having homologs already mapped in lettuce and sunflower, thereby facilitating comparative analyses across the family. The CGP will be also involved in the sequencing of globe artichoke gene space (i.e., the low copy, gene-rich fraction of the genome) using Illumina sequencing technology, enhancing genome-level structural information, exploitable in agricultural key traits dissection.
      Publications
    Acquadro, A., Lanteri, S., Scaglione, D., Arens, P., Vosman, B. and Portis, E. (2009) Genetic mapping and annotation of genomic microsatellites isolated from globe artichoke. Theoretical and Applied Genetics, 1573-1587.

    Basnitzki, J. and Zohary, D. (1994) Breeding of seed planted artichoke. Plant Breeding Reviews 12, 253-269.

    Curt, M., Sanchez, G. and Fernandez, J. (2002) The potential of Cynara cardunculus L. for seed oil production in a perennial cultivation system. Biomass & Bioenergy, 33-46.

    Encinar, J., Gonzalez, J. and Gonzalez, J. (2002a) Steam gasification of Cynara cardunculus L.: influence of variables. Fuel Processing Technology, 27-43.

    Encinar, J., Gonzalez, J., Rodriguez, J. and Tejedor, A. (2002b) Biodiesel fuels from vegetable oils: Transesterification of Cynara cardunculus L. oils with ethanol. Energy & Fuels, 443-450.

    Foti, S., Mauromicale, G., Raccuia, S., Fallico, B., Fanella, F. and Maccarone, E. (1999) Possible alternative utilization of Cynara spp. I. Biomass, grain yield and chemical composition of grain. Industrial Crops and Products, 219-228.

    Foury, C., Martin, F., Vaissière, B., Morison, N., and Corre, J., 2005, Advantages et Difficultes de la Creation d’Hybrides F1 d’Artichaut à Semer, Acta Horticulturae. 681:315-322.

    Ierna, A. and Mauromicale, G. (2010) Cynara cardunculus L. genotypes as a crop for energy purposes in a Mediterranean environment. Biomass & Bioenergy, 754-760.

    Lanteri, S., Acquadro, A., Comino, C., Mauro, R., Mauromicale, G. and Portis, E. (2006) A first linkage map of globe artichoke (Cynara cardunculus var. scolymus L.) based on AFLP, S-SAP, M-AFLP and microsatellite markers. Theoretical And Applied Genetics 112, 1532-1542.

    Lanteri, S. and Portis, E. (2008) Globe Artichoke and Cardoon. In: Vegetables I (Springer ed) pp. 49-74. New York: Springer.

    Lapuerta, M., Armas, O., Ballesteros, R. and Fernandez, J. (2005) Diesel emissions from biofuels derived from Spanish potential vegetable oils. Fuel, 773-780.

    Lattanzio, V., Kroon, P.A., Linsalata, V. and Cardinali, A. (2009) Globe artichoke: A functional food and source of nutraceutical ingredients. Journal of Functional Foods 1, 131-144.

    Lombardo, S., Pandino, G., Mauromicale, G., Knödler, M., Carle, R. and Schieber, M. (2010) Influence of genotype, harvest time and plant part on polyphenolic composition of globe artichoke [scolymus (L.) Fiori]. Food Chemistry 119, 1175-1181.

    Mauro, R., Portis, E., Acquadro, A., Lombardo, S., Mauromicale, G. and Lanteri, S. (2009) Genetic diversity of globe artichoke landraces from Sicilian small-holdings: implications for evolution and domestication of the species. Conservation Genetics: 10:431-440.

    Mauromicale, G. and Ierna, A. (2000) Panorama varietale e miglioramento genetico del carciofo. Informatore agrario 26, 39-45.

    Pandino, G., Courts, F., Lombardo, S., Mauromicale, G. and Williamson, G. (2010) Caffeoylquinic Acids and Flavonoids in the Immature Inflorescence of Globe Artichoke, Wild Cardoon, and Cultivated Cardoon. Journal of Agricultural and Food Chemistry, 1026-1031.

    Pandino, G., Lombardo, S., Mauromicale, G. and Williamson, G. (2011) Phenolic acids and flavonoids in leaf and floral stem of cultivated and wild Cynara cardunculus L. genotypes. Food Chemistry, 417-422.

    Pignone, D. and Sonnante, G. (2004) Wild artichokes of south Italy: did the story begin here? Genetic Resources and Crop Evolution 51, 577-580.

    Portis, E., Barchi, L., Acquadro, A., Macua, J. and Lanteri, S. (2005b) Genetic diversity assessment in cultivated cardoon by AFLP (amplified fragment length polymorphism) and microsatellite markers. PLANT BREEDING 124, 299-304.

    Portis, E., Mauromicale, G., Barchi, L., Mauro, R. and Lanteri, S. (2005a) Population structure and genetic variation in autochthonous globe artichoke germplasm from Sicily Island. Plant Science 168, 1591-1598.

    Portis, E., Mauromicale, G., Mauro, R., Acquadro, A., Scaglione, D. and Lanteri, S. (2009) Construction of a reference molecular linkage map of globe artichoke (Cynara cardunculus var. scolymus). Theoretical and Applied Genetics, 59-70.

    Rottenberg, A. and Zohary, D. (1996) The wild ancestry of the cultivated artichoke. Genetic Resources And Crop Evolution 43, 53-58.

    Scaglione, D., Acquadro, A., Portis, E., Taylor, C., Lanteri, S. and Knapp, S. (2009) Ontology and diversity of transcript-associated microsatellites mined from a globe artichoke EST database. BMC Genomics, 10: 454.

    Schutz, K., Kammerer, D., Carle, R. and Schieber, A. (2004) Identification and quantification of caffeoylquinic acids and flavonolds from artichoke (Cynara scolymus L.) heads, juice, and pomace by HPLC-DAD-ESI/MSn. Journal Of Agricultural And Food Chemistry 52, 4090-4096.
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