Crop Case Study Beta L. (Including Patellifolia AJ Scott Et Al.)
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Crop case study Beta L. (including Patellifolia A. J. Scott et al.) L. Frese, M.A Pinheiro de Carvalho, C. Duarte Introduction Beta vulgaris L. subsp. maritima Arcang. is considered the ancestor of all cultivated beets. Zossimovich (1939) cited by Boughey (1981) assumed that the cultivated beets were first grown on irrigated fields of ancient Mesopotamia but fossiles supporting this theory were never excavated. There is however historical evidence that the cultivation of beets as green diet was well known to the Greek people in 370 B.C. At that time Theophrastus even described the differences between wild and cultivated forms (Boughey, 1981). Beets cultivated as green salad vegetable are likely the ancestor of garden, fodder and sugar beets, the development of the latter started only 200 years ago. From an evolutionary point of view the sugar beet was developed within a very short time towards a high productive crop. Bosemark (1989) suggested therefore maintaining the adaptability of sugar beet by continuous and systematic incorporation of genetic variability from the closest relative, i.e. Beta vulgaris subsp. maritima, into the elite gene pool. Biancardi (2005) reviewed the objectives of sugar beet breeding and devoted several chapters to resistance breeding. Genetic protection of the sugar beet crop against viruses (Curly Top, Beet Yellows, Beet mosaic, Rhizomania, Beet soil borne virus), bacteria (Bacterial vascular necrosis and rot, Yellow wilt), fungi (Cercospora leaf spot, Powdery mildew, Downey mildew, Fusarium yellows, Root rots such as Rhizoctonia, Phoma, Pythium, Aphanomyces, Sclerotium), Polymyxa, Alternaria, Ramularia, Uromyces), and pests (Cyst and root knot nematode, Aphids, Spinach leaf miner, Sugar beet root maggot, Flea beetle and last not least Spider mites are welcomed by growers and consumer since it helps to keep production costs lower and helps reducing pollution of the environment with fungicides and insecticide residues. To achieve the resistance breeding objectives breeders have used wild beets as a source of novel genes. Steinrücken (2005) anticipates an even growing demand for genetic resources. He argues that the improved breeding techniques based on molecular marker technologies and an improved knowledge of the Beta genome will facilitate the integration of genes from exotic germplasm into the crop and therefore increase the interest in Beta genetic resources and the demand for novel genetic variation. As a consequence of climate warming growing conditions are changing at accelerated rates and from the evolutionary perspective within an extremely short period. The domesticated species developed from their highly diverse wild ancestors may not always have the genetic variation available needed to cope with these fast changes. The genetic reserve conservation technique (Maxted et al., 1997) recognizes the need for maintenance of the intraspecific diversity which is the natural basis of adaptation of any species to environmental changes and a crucial condition for its survival in rapidly changing environments. A wide genetic basis of wild species related with a crop ensures the evolution of new genes and alleles required for the adaptation of that crop. There are therefore strong arguments to protect crop wild relatives in the natural surrounding where they have developed their adaptive traits as well as traits currently used by plant breeders. The establishment of genetic reserves is thus a service of nature and species conservation to agriculture. The design, planning and establishment of genetic reserves are technical conditions for the maintenance of intraspecific diversity. A step-wise methodology for the identification of genetic reserve sites for a target crop gene pool was developed by the University of Birmingham within the AEGRO project. The methodology can be accessed online at http://aegro.bafz.de/index.php?id=188. Recommendations for the choice of genetic reserves sites for Beta and Patellifolia (syn. Beta section Procumbentes Ulbrich) within the EU-27 as well as in Portugal and Germany described in the case study are based on this 4-step-methodology. Application of the four step methodology The case study consists of three parts. As there is no need to repeat step 1 of the methodology for the national level case study, the case study begin with the taxon delineation and continues with EU-27 level case study (Part A) followed by the national level case study (Part B). Step 1: Taxon delineation Information on the distribution of wild species can be derived from various information systems and data sources. Since there is no single globally accepted taxonomic system for Beta three systems were implemented in the CWRIS AEGRO Population Level Information System (PLIS) (CWRIS AEGRO PLIS, 2010). The case study is based on a taxonomic system proposed by Frese. The relation of sections, species and gene pools is presented in Table 1. The fourth section of the genus (Beta section Procumbentes) has been accepted as the new genus Patellifolia (Scott et al., 1977). Although Patellifolia species are no longer part of Beta they should be treated as wild beets belonging to the gene pool of cultivated beets. This suggestion is well in agreement with the definition of a gene pool published by Harlan and de Wet (1971). The suggestion of Kadereit et al. (2006) to include B. nana Boiss. & Heldr. into Beta section Corollinae Ulbich was also considered by the working taxonomy for Beta shown in Table 1. Table 1. Beta working taxonomy and assignment of taxa to the primary, secondary and tertiary gene pool of cultivated beets Gene pool Genus Beta L. Primary Section Beta Transhel B. vulgaris L. subsp. vulgaris (cultivated beets) Leaf Beet Group Garden Beet Group Fodder Beet Group Sugar Beet Group subsp. maritima (L.) Arcang. subsp. adanensis (Pamuk.) Ford-Lloyd & Will. B. patula Ait. B. macrocarpa Guss. Secondary Section Corollinae Ulbrich Base species B. corolliflora Zosimovich B. macrorhiza Steven B. lomatogona Fisch & Meyer Hybrid species B. intermedia Bunge B. trigyna Wald. & Kid. B. nana Boiss. & Heldr. Tertiary Genus Patellifolia A. J. Scott et al. P. procumbens (Smith) A.J.Scott, Ford-Lloyd & J.T.Williams P. webbiana (Moq.) A.J.Scott, Ford-Lloyd & J.T.Williams P. patellaris (Moq.) A.J.Scott, Ford-Lloyd & J.T.Williams Although the identification of genetic reserve sites for CWR is the main purpose of this case study the taxonomy of cultivated beets is commented here. A future study should consider cultivated forms to facilitate decision to be taken in the field of on farm management. The highly detailed taxonomy of cultivated beets used by many gene banks (“splitter system”) requires intimate knowledge of the meaning of taxonomic names and cannot be used straightforward by sporadic users of information systems such as Global Biodiversity Information Facility (GBIF). In particular the downloaded GBIF file contained a large number and variation of taxonomic names of cultivated forms which have been processed and harmonized along with the wild species taxonomic names. The taxonomic names used by “splitter systems” were matched to the “lumber system” for cultivated types. The latter was proposed by Lange et al. (1999) who have sorted all cultivated forms into four cultivar groups. This simple and clear classification proposed by Lange et al. (1999) is part of the Frese system used by CWRIS AEGRO PLIS but has a disadvantage: A highly detailed splitter system can be mapped to the lumber system. Mapping in the opposite direction is not possible as long as the accessions classified according to Lange et al. (1999) has not been characterised in detail. An 2 example: Beta vulgaris subsp. vulgaris convar. cicla var. flavescens forma rhodopleura is classified by Lange et al. (1999) as Beta vulgaris subsp. vulgaris culton Leaf Beet group. The “convar. cicla var. flavescens forma rhodopleura” describes a red coloured leaf beet with, while “culton Leaf Beet groups” circumscribes a leaf beet, only. Not all relationships between taxonomic names used to describe variation in the cultivated species have been elaborated during the preparation of the information basis. In particular the Russian informal taxonomic system has not yet been analysed in detail and mapped to the three systems implemented in CWRIS AEGRO PLIS since work on cultonomy is not part of this case study. A list of synonyms for taxonomic names contained in information systems and/or described in literature can be found at http://aegro.bafz.de/aegroprod_beta/gm/queryBuilder/findTaxon.seam. Part A of the case study: EU-27 Step 2: Selection of target taxa According to Ford-Lloyd et al. (2009) genetic reserves should be established if a taxon occurs is less than 10 geographic units. Geographic units are listed in the Crop Wild Relative Information System (CWRIS, 2009). This concept was applied and yielded the following list. Taxa printed in bold letter are naturally distributed within the EU-27 while the other species may be encountered due to sporadic carryover into EU-27 countries. B. patula B. macrocarpa (4x) B. vulgaris subsp. adanensis B. corolliflora B. macrorhiza B. lomatogona B. nana B. procumbens Smith (=Patellifolia procumbens) B. webbiana Moq. (=Patellifolia webbiana) B. patula is confined to two islets off the main island Madeira (Pinheiro et al., 2010) which counts for a single geographic unit. B. macrocarpa occurs in more than 10 geographic units. Until recently it was assumed that the diploid form is mainly distributed in Portugal, Spain, Greece, Israel and Morocco while the distribution of the tetraploid form is restricted to the Canary Islands. This assumption was revised by Villain (2007) who detected diploid B. macrocarpa on the Canary Islands. Tetraploid forms have not been found in the continental part of the distribution area to date as allopolyploid offsprings of B. vulgaris subsp. maritima x B. macrocarpa are extremely rare in the Iberian part of the distribution area. Villain (2007) confirmed Lange and de Bock (1989) who had suggested an allopolyploid origin of the tetraploid type of B.