sign in sign up
<<
Home , Brassica, Brassicaceae, Brassica carinata, Brassica oleracea, Brassica rapa, Brassica tournefortii

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 www.inacj.com ISSN: 2228-6322© International Academic Journals

The wild and the grown – remarks on

Hammer K.¹*, Gladis Th.¹ , Laghetti G.² , Pignone D.² ¹Former Institute of Crop Science, University of Kassel, Witzenhausen, . * Author for correspondence (email: [email protected]) ²CNR – Institute of Genetics, Bari, .

Received mmmm yyyy; accepted in revised form mmmmm yyyy

ABSTRACT Brassica is a of the Cruciferae (). The wild races are concentrated in the Mediterranean area with one in CE (Brassica somaliensis Hedge et A. Miller) and several weedy races reaching E . Amphidiploid evolution is characteristic for the genus. The diploid species (L.) Koch (n = 8), L. emend. Metzg. (n = 10, syn.: B. campestris L.) and L. (n = 9) all show a rich variation under domestication. From the naturally occurring amphidiploids (L.) Czern. (n = 18), Brassica napus L. emend. Metzg. (n = 19) and the rare A. Braun (n = 17) also some races have developed. The man-made Brassica ×harmsiana O.E. Schulz (Brassica oleracea × Brassica rapa, n = 29, n = 39), or similar hybrids, serve also for the development of new . Gouan (n = 10) from another Brassica- cytodeme, different from the Brassica rapa group, is occasionally grown as a vegetable in . Brassica has developed two hotspots under cultivation, in the Mediterranean area and in E Asia. Cultivation by man has changed the different Brassica species in a characteristic way. The large amount of morphologic variation, which exceeded in many cases variations occurring in distinct wild species, has been observed by the classical botanists by adding these variations to their natural species by using Greek letters. Later taxonomists used the category botanical variety (var.). In this way impressive systems have been established, e.g. for Brassica oleracea. Later on, the other species followed. The variation from E Asia, particularly in the species Brassica rapa and Brassica juncea, was much later included into the investigations, simply because of lacking information. However, this material was included, in the last one hundred years, in classifications according to the International Code of Botanical Nomenclature, ICBN (McNeill et al. 2006). An overview is provided of the infraspecific taxa in Brassica vegetable species. This is one possibility to demonstrate the rich variation of the cultivated races. Included here are our experiences with field studies in the Mediterranean area (especially Italy) and E Asia (Korea, ). A short discussion is devoted to the classification of material according to the International Code of Nomenclature for Cultivated , ICNCP (Brickell et al. 2009). Reducing the possibilities for classification largely to and groups, this Code is certainly adapted to modern agri- and , including the seed business. Whereas cultivated plants experience a fast evolution, depending on the speed of breeding progress in different periods of the last less than 10.000 years, starting from the so called Neolithic revolution, wild plants are following usually slower evolutionary pathways over a much longer time span. It is difficult to classify wild and cultivated plants under the same system. The genus Brassica provides material, also showing a complex reticulate evolution difficult to classify, which can help developing ways for a general framework.

Keywords: Brassica spp., vegetable, classification, genetic resources, wild plants, weedy plants, cultivated plants

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 453

INTRODUCTION Oilseed include Brassica carinata (cultivated in Ethiopia and North-east Brassica is widely distributed in the Africa), B. napus (cultivated in and Mediterranean with about 20 species. The ) and B. juncea (mainly scope of the genus is still under discussion, cultivated in South- and South-). with some researchers including into They are amphidiploids combining genomes Brassica also L. (with S. alba L. of the diploid species B. rapa , B. nigra and [Brassica hirta Moench], an important oil B. oleracea as already found out by U (U seed plant and also vegetable in the 1935) who is well known for the triangle of Mediterranean and the farmweed S. arvensis U. Several Brassica oil seeds, including B. L. [Brassica kaber (DC.) Wheeler], see carinata, B. juncea , B. napus and the diploid Baillargeon 1986). Also other genera show B. rapa, naturally produce seed oil moderate close relationships as evidenced also by to high in (22:1cis13) content and crossing experiments. The genus moderate to high content in the seed L. has produced hybrids with Brassica of meal after oil extraction (Downey and some economic importance Röbbelen 1989). Ranges of erucic acid (× Sageret, syn. content in these species have been reported ×Raphanobrassica Karpechenko, see Specht by Velasco et al. (1998) as: B. carinata: 2001). The genus Raphanus is possibly 29.6-51.0%; B. juncea: 15.5-52.3%; B. derived from B. nigra and B. rapa/oleracea napus: 5.6-58.1% and B. rapa: 6.5-61.5%. hybrids (Mabberley 2008). Mill. is Black and also naturally another close relative and possible candidate produce seed oil with a range in erucic acid for inclusion (Pignone and Gómez Campo content. Tahoun et al. (1999) have reported 2010). Modern taxonomists speak about the erucic acid content ranges for B. nigra: 30.3- --Brassica complex 45.0% and for B. oleracea: 0.1-62.0%. The (Sánchez-Yélamo 2009). plants and seeds of all Brassica oilseeds The free crossability of closely related contain , secondary wild/weedy/ and cultivated races sharing the metabolites which serve as chemical same genome led to proposals for lumping, protectants (Mitten 1992). Ranges of e.g. for the Brassica oleracea group (Gladis content in these species have and Hammer 2001), with reduction on the been reported to be 20 to > 200 µmol g-1 number of species. seed total glucosinolates for B. napus, B. Some species of Brassica (e.g. B. rapa, oleracea and B. rapa; 75 to > 150 µmol g-1 Zohary et al. 2012) have been early in seed total glucosinolates for B. carinata and history used for their seeds as an oil crop. B. nigra and 100 to > 200 µmol g-1 seed total Today, the development of some Brassica glucosinolates for B. juncea (Röbbelen and species towards a crop for edible and Theis 1980). industrial oils uses has reached a great Different Brassica oilseeds species importance (Daun et al. 2011). They are the predominate in different regions of the third most important source of vegetable oils world. In the warmer semi-tropical regions, after oil palm and soybean (Gupta and B. juncea and B. rapa predominate while in Pratap 2007). Oilseed brassicas have been cooler temperate regions B. napus and B. recently treated in more detail (Diederichsen rapa. B. carinata is limited to Ethiopia and and McVetty 2011). Therefore, only some northeast Africa, while B. nigra is grown in results are summarized in this introduction. Europe and Asia. B. nigra is currently grown

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 454

mainly as a condiment crop. B. oleracea is Winter B. rapa types, formerly grown in exclusively a vegetable crop produced northern Europe, have been replaced by globally. B. juncea is an important oilseed higher yielding winter B. napus types. B. species in Asia and also an important tournefortii is cultivated on a small scale in condiment crop in that has recently India (Diederichsen 2001). been converted to a new Canadian edible Here, the vegetable brassicas will be treated oilseed crop (Potts et al., 2003). B. napus is in more detail. They include after the predominate oilseed species in , Diederichsen (2001) Brassica oleracea, B. Europe, Canada, and China, while B. juncea rapa, B. juncea, B. napus, B. ×harmsiana, B. is the predominate species in India and carinata, and B. nigra. Gladis and Hammer northwest China. Winter B. napus types are (1990) also mention Brassica tournefortii. grown in southern Europe while spring B. There is a high range of different usages, napus and B. rapa types are grown in which also include different plant parts (see northern Europe (Gupta and Pratap, 2009). table 1).

Table 1. Cultivated brassicas in the different usage groups indicating also the plant parts used(after Gladis and Hammer 1990, modified) Organ / Brassica species 1 2 3 4 5 6 7 8 and Hypocotyls -- -- V FV -- -- VF -- Shoots and V F GVD FGV GFV VFD VFGD V -- -- V V -- V V -- B B BV BV B VB BV -- Seeds OM O OMFG OF OMG -- OF O 1 = Brassica carinata, 2 = B. ×harmsiana, 3 = B. juncea, 4 = B. napus, 5 = B. nigra, 6 = B. oleracea, 7 = B. rapa, 8 = B. tournefortii Usage groups (ordered according to their importance): B = bee-, G = green manuring, F = fodder, V = vegetable, O = oilseed, M = medicinal and spice plant, D = decorative and ornamental plant, -- = not known

There are experiences from field studies of to be included. The book of Ambrosoli the authors of this paper from the primary (1997), who studied the fodder plants, centre of variation of Brassica in the relatively new crops with first origins in the Mediterranean area, especially from Italy classical world and a later development in N (Hammer et al. 1992, 1999, 2011) and in E Italy and W Europe (1350 – 1850). Their Asia (Hoang et al. 1997, Hammer et al. and , has been taken as a 2007, Li et al. 2011). During several model for our treatment, as can be seen also collecting missions, which are documented from the title of our paper. in the compilations cited above, emphasis was laid on cultivated plants, crop wild MORPHOLOGICAL DIVERSITY OF relatives (Maxted et al. 2008) played an BRASSICA SPECIES important role from early phases of our A brief overview over morphologically exploration (e.g. Perrino et al. 1992). based classifications of cultivated Brassica As many Brassica spp. contain cultivated, vegetable species and their wild relatives is wild and weedy races, insights into provided here. The most important species evolutionary pathways were possible within are treated, including also the races used as this genus of traditional vegetable crops, oil crops. A wide range of geographic which surely belong to the oldest crop distribution and a tremendous range of plants. Elements of human behavior should different usages lead to an enormous be included in this survey. Together with phenotypic range of diversity in Brassica. botanical, genetical and horticultural Homologous developments for usages and elements, a broad scope of information has organs used occurred in different species

455 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

(see table 1). As for several usage groups ICNCP it can still be found. Fortunately we and crop types, common names are not can rely on ICBN and its infraspecific available in English, we prefer the Latin possibilities (except convar. which was names according to ICBN (McNeill et al. under ICNCP). In Brassica since Linnaeus 2006). Naming according to the ICNCP (1753, 1763) a formal infraspecific (Brickell et al. 2009) is not preferable in classification has been created and further Brassica because of the limited possibilities developed by De Candolle (1824), Alefeld for infraspecific classification (only cultivar (Alefeld 1866) and others. Formal and cultivar groups), which may be classification proved to be useful in sufficient for growers and seed business, but monitoring the decline of diversity in not for researchers engaged in and agriculture (Vellvé 1993, Hammer et al. evolution. The lacking of handling of 1996). Contrary to the trend to use informal landraces according to ICNCP has already classifications (Toxopeus and Oost 1985, criticized by Pickersgill and Karamura see Hanelt 1986 b), for an example see table (1999). Another change in the development 2), formal classifications for Brassica are of the ICNCP is the removing of the very still in predominant use ( some recent useful category of convar. ( is examples are: Louarn et al. 2007, Qi et al. no substitute), which has been proposed by 2008, van Treuren and Bas 2008, Wu et al. Alefeld (1866, see also Landsrath and 2009, Cleemput and Becker 2012, Girke et Hammer 2007) and was advocated e.g. by al. 2012). Mansfeld (1959). In the first issues of

Table 2. Examples comparing informal and formal classification of Brassica rapa (after Diederichsen 2001, modified) Utilized plant organ(s) Informal group Formal classification Vegetable group ssp. rapa Root Fodder turnip group ssp. rapa Root and Leaf Leaf turnip ssp. chinensis var. chinensis Leaf Chinese ssp. pekinensis var. glabra Leaf Pak Choi and Saishin ssp. chinensis var. chinensis et var. purpuraria Leaf Taatsai ssp. chinensis var. rosularis Leaf ssp. nipposinica and young leaves Broccoletto ssp. oleifera var. ruvo Seed Winter turnip rape ssp. oleifera var. oleifera f. autumnalis Seed Spring turnip rape ssp. oleifera var. oleifera f. praecox Seed Brown sarson and toria ssp. dichotoma Seed sarson ssp. trilocularis var. trilocularis et var. quadrivalvis

Brassica oleracea L. Brassica crops and inspired a lot of research Brassica oleracea has no distinct race for and breeding activities in the genus Brassica seed use, but is very rich in diversity for (Snowdon 2007). One of the diploid species vegetable, forage, ornamental and other is B. oleracea (n=9) which has been uses. The genetic relationships among the cultivated for a long time. The wild cultivated taxa of the genus Brassica were progenitors of the different cultivated B. studied by various cytologists in the oleracea races are the perennial subspecies beginning of the 20 century and in 1935 U of B. oleracea (as proposed by Gladis and summarized the results by presenting what is Hammer 2001). Yarnell (1956), Snogerup frequently referred to as U’s triangle (U (1980) and others have proposed that 1935, Helm 1963, Mizushima 1980). U’s different taxa of this group were involved in concept has been extremely fruitful for the evolution of the various vegetable races understanding the evolution of the important of the cultivated taxa of B. oleracea. The

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 456

closest wild relative is the wild perennial populations. The Potuguese “Galega” kales cabbage of the European Atlantic coast, B. (Dias 1995) may belong here. oleracea subsp. oleracea (Snogerup et al. ssp. (L.) DC. s.l., here, all cultivated 1980, Song et al. 1990, Hodgkin 1995, are included (Gladis and Hammer Gustafsson and Lannér-Herrera 1997). 2003). Glucosinolate investigations support a convar. fruticosa (Metzg.) Alef. Most multiphyletic origin for cultivated forms primitive and perennial types with wooden from different races (subspecies) of wild and branched shoots. Only one variety Brassica (Mithen et al. 1987). On the basis present. of linguistic, literary and historical var. ramosa DC. perpetual kale. A primitive information, Maggioni et al. (2010) variety with ramifications (cottager’s kale). . postulated for the origin of cultivated races Leaves used as a vegetable or of Brassica oleracea the ancient Greek fodder. This perennial kale today can be speaking areas of the Mediterranean, namely found as a relic crop in W Europe (Zeven et Magna Grecia. (c. 372 – 287 al. 1998, Dias 2012) and some tropical areas BC) already described different races of (Diederichsen 2001). Brassica oleracea, e.g.with curly and smooth convar. acephala (DC.) Alef. All not leaves (Hondelmann 2002). branched vegetable races for leaf use are Brassica oleracea is a good example where included here. As already discussed, the wild and cultivated races show clearly category convar. has been excluded from the differing pathways. The wild races, mainly latest version of ICNCP (Brickell et al. from the Mediterranean, reached several 2009). However, this category is important islands, mainly swimming and thus showing for variable cultivated plants and, therefore large fruits. They are adapted to calcareous it is maintained here traditionally (see Helm cliffs where they could successfully escape 1960 – 1963). goats and fire. Their populations on less var. acephala DC. (excl. var. ramosa). protected grounds have already disappeared Primitive races from southern Italy may long ago and only recently a reduced belong here (Branca and Iapichino 1997, selection pressure brings them back in the Branca et al. 2010) and also the above neighbourhood of gardens where mentioned “Galega” kales. introgressions with the cultivated races var. gongylodes L. . Helm (1959) could be observed (Perrino and Hammer lists three different forms: f. gongylodes (L.) 1985). From the wild races ssp. rupestris, Helm (with green stem tubers), f. violacea ssp. insularis, ssp. macrocarpa and ssp. Lam. (with violet stem tubers), f. dissecta villosa formed a cluster, genomically not Peterm.(with laciniate leaves). A similar closely related to the cultivated types (Mei race was described by A.P. De Candolle as ß et al. 2010). The wild races of the Brassica crispa DC., Syst. Nat. 2 (1821) 586, based oleracea cytodeme can be seen from table 3 . on the Italian landrace “Pavonazza” which Cultivated races was grown around Naples (Helm 1963). A ssp. capitatoides Gladis et K. Hammer, nom. similar race was found by us in Apulia prov. The plants are derived from crosses (Hammer et al. 1989, fig. 10). Primitive between wild and cultivated B. oleracea. kohlrabis with large tubers (Germ. Strunk- They are usually stabilized crossing Kohlrabi) are relic crops in some countries, products (Snogerup 1980, Gladis and e.g. “kana pchali” in Georgia (Beridze et al. Hammer 2003). Plants mostly semi- 1983). cultivated, sometimes wild, forming small

457 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Table 3. The wild races of the Brassica oleracea cytodeme Species Number and Infraspecific Classification Status caryotype Brassica oleracea L. n=9, C ssp. oleracea Wild n=9, C ssp. bourgaei (Webb in Christ) Gladis et K. Hammer Wild n=9, C ssp. cretica (Lam.) Gladis et K. Hammer Wild var. aegaea (Heldr. et Hal.) Gladis et K. Hammer var. cretica (Lam.) Coss. var. laconica (Gust. et Snog.) Gladis et K. Hammer var. nivea (Boiss. et Sprun.) Gladis et K. Hammer n=9 ssp. hilarionis (Post) Gladis et K. Hammer Wild n=9 ssp. incana (Ten.) Gladis et K. Hammer Wild var. botteri (Vis.) Gladis et K. Hammer var. cazzae (Ginzberger et Teyber) Gladis et K. Hammer var. incana (Ten.) Gladis et K. Hammer var. mollis (Vis.) Gladis et K. Hammer n=9 ssp. insularis (Moris) Rouy et Fouc. Wild var. angustiloba Schulz var. aquellae Widler et Bouquet var. atlantica (Coss.) Batt., Batt. et Trab. var. conferta (Jordan) Schulz var. insularis (Moris) Coss. var. latiloba Schulz n=9 ssp. macrocarpa (Guss.) Gladis et K.Hammer Wild var. drepanensis (Caruel) Schulz var. macrocarpa (Guss.) Gladis et K. Hammer n=9 ssp. robertiana (Gay) Gladis et K.Hammer Wild n=9 ssp. rupestris (Raf.) Gladis et K.Hammer Wild var. brevisiliqua (Raimondo et Mazzola) Gladis et K. Hammer var. hispida (Raimondo et Mazzola) Gladis et K. Hammer var. rupestris (Raf.) Paol. n=9 ssp. villosa (Bivona-Bernardi) Gladis et K.Hammer Wild var. bivoniana (Mazzola et Raimondo) Gladis et K. Hammer var. taurica (Tzvel.) Gladis et K. Hammer var. tinei (Lojac.) Gladis et K. Hammer var. villosa (Biv.) Coss. For further brassicas and wild/occasionally cultivated relatives see Diederichsen and McVetty 2011 var. medullosa Thell. marrow-stem kale. Recently, a long-stalked, purple leaved Mainly used as fodder plant. Possibly variety of the latter is commonly used as originated from a cross of var. gongylodes ornamental plant in Germany. and var. viridis (Gladis and Hammer 2003). var. selenisia L. feather kale. Formerly a Already mentioned by De Candolle (1824) common garden plant, today rather rare. as “chou moellier”(Helm 1959). Helm (1963) differentiates f. selenisia Helm var. palmifolia DC. palm kale. An old race. with green and f. scotica (Alef.) Helm with Today still widely grown as delicious purple or partly coloured foliage. Races vegetable in Italy (Hammer et al. 1990). with feathery and laciniated leaves occur var. sabellica L. curled kitchen kale. Still also in other botanical varieties of B. wide spread as a cooking vegetable with oleracea (vgl. Helm 1962). good winter hardiness. Often united with the var. viridis L. kale, . Occasionally following variety (e.g. by Helm 1959). Helm grown for fodder, young leaves for human (1963) described f. sabellica (L.) Helm consumption. Helm (1959) describes three (green) and f. rubra Peterm. (purple-violet). races: f. viridis L. (green), f. purpurascens

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 458

(DC.) Thell. (red) and f. exaltata (Rchb.) (Branca and Iapichino 1997). Thell., Jersey kale, the long, lignified stems Helm (1963) described f. erytrobotrys of which are used for roof constructions and (Alef.) Helm (violet), f. phaeusa (Alef.) walking sticks (Hew and Rumball 2000). Helm (brown), f. chlorusa (Alef.) Helm The first two forms are very traditional in (green), f. theiusa (Alef.) Helm (yellow) and Italy. Also relic crop in the US (Farnham et f. botrytis (L.) Helm (White). After centuries al. 2008). with predominant white inflorescences, convar. botrytis (L.) Alef. This group unites more and more varieties with coloured heads all races which are grown for the use of (yellow, green, purple) can be found on the young inflorescenses. ”Mugnoli” from S markets today. Italy can be seen as a primitive race in this var. alboglabra (Bail.) Sun ex Musil. group. The fleshy deformation of the Chinese kale. Evidently originated in the inflorescenses is just developing in this race Mediterranean area. Similar races have (Laghetti et al. 2005). This may be a relic of been collected in S Italy (Maly et al. 1987) introductions from eastern areas (e.g. Crete) and S Spain (Hammer in Gladis and together with Greek (Laghetti et al. 2008) or Hammer 2003). The progenitors probably Albanian (Hammer et al. 2012) immigrants have been moved along the silk road and with which the evolution of var. italica and reached E Asia several centuries ago. Under var. botrytis started. In the Salento of the specific selection conditions of China Apulia this race can still be found. The this speciality arose. Crisp and Gray (1984) glucosinolate profile of “Mugnoli” turned placed it to the . Helm (1964) and out to be closer to var. italica (Argentieri et Larkcom (1987) gave detailed descriptions. al. 2010). Variable material of convar. A fertile amphidiploid with botrytis was found in Sicily (Branca and Raphanus sativus was produced (Chen and Iapichino 1997). Wu 2008). var. italica Plenck. . From convar. capitata (L.) Alef. s.l. This is the an Italian speciality it has developed in the group of the common cabbages. They last decennia to a world crop. Purple and originated in the Mediterranean area and green heading broccolis are recently placed have a wide distribution. to this variety. As already stated by var. capitata L. common cabbage. With Vilmorin (Helm 1959), broccoli is older different colours and head-forms (Helm than cauliflower. As areas of origin Helm 1963). Economically most important race of (1959) mentioned South Italy, Cyprus and B. oleracea. Eaten as a vegetable raw or Crete. South Italy is still rich in landraces. cooked. Conserved for later use by Helm (1963) described f. cymosa Duch. fermentation as . After Helm (green), f. albida Duch. (white), f. flava (1963) with f. subacuta Duch., f. conica Peterm. (yellow) and f. italica (Plenck) Duch. and f. capitata (L.) Helm. Helm (violet). var. sabauda L. . Derived var. botrytis L. cauliflower. Developed in from a cross of var. capitata and var. the Mediterranean region and became a palmifolia (Gladis 1995). Eaten as a world crop. Related to Brassica oleracea ssp. vegetable cooked and raw, especially in cretica. The introduction from Crete to winter. Possibly originated in Italy (Helm South Italy could have happened by 1959). Helm (1963) described f. ovata immigrants from the Levante (Hammer et al. Duch., f. oblonga (DC.) Peterm. and f. 2011). Sicily and the Naples area played an sabauda (L.) Helm. For the classification of important role in the further evolution of the eastern cabbages see also Lizgounova in

459 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Zhukovsky (1933). Vilmorin-Andrieux sylvestris) has ( possibly extinct?) its natural presented races with laciniated leaves which distribution (now mostly weedy races, but obviously have not be maintained (Gladis Andersen et al.( 2009) discuss about wild and Hammer 2003). races in parts of N Europe) and East Asia convar. costata (DC.) Gladis ex with (only?) weedy types of secondary Diederichsen nature. The result in the areas is var. costata DC. tronchuda kale. The predominant seed use and vegetable use, thickened leaf stalks and leaf ribs, respectively (Sinskaja 1928, Warwick et al. sometimes the leaf blades are used as a 2008). Chinese cabbages originated in cooking vegetable. Formerly grown in China more than 6.000 years ago. Seed oil western Europe, now mainly in , types came from northern Europe to China Angola and . and Korea about 2.000 years ago. Two var. helmii Gladis et K. Hammer. Derived independent domestication events in Europe from a cross of var. capitata and var. (Reiner et al. 1995) and E Asia have been sabellica. Today mostly grown as an postulated by McGrath and Quiros (1992). ornamental plant (Gladis and Hammer According to Takuno et al. (2007) a further 2003). One old German variety with domestication in E Asia occurred after yellowish-green foliage named ‘Mosbacher introduction of primtive cultivated types Winter Hellgrüner Blätterkohl’ had been from Europe and Central Asia. Later, the used as vegetable. diploid Brassica rapa hybridized convar. gemmifera (DC.) Gladis ex spontaneously with Brassica oleracea and Diederichsen. Races with multiple leaf buds Brassica nigra in areas where their on one stalk. geographical ranges of distribution under var. polycephala Thell. (syn. var. cultivation overlapped resulting in three delachampii Helm (excl. var. millecapitata amphidiploid bastards, B. juncea (n=18), B. (Lév.) Helm; after Thellung with open carinata (n=17) and B. napus (n=19), that rosettes of leaves, therefore syn. of var. are cultivated species of their own, with ramosa!), for nomenclatural details see occasional weedy derivates. The origin of Helm 1959). A race with few multiple the Indian races, which are grown as heads. Formerly a common garden plant. oilseeds, is still under discussion. Possibly Perhaps ancestor of var. gemmifera. ssp. dichotoma and ssp. trilocularis belong var. gemmifera DC. Brussels sprouts. The to the western Eurasian branch together with axillary sproutlets are used as vegetable. ssp. rapa and ssp. oleifera, whereas ssp. Very successful race. Today distributed all chinensis, ssp. pekinensis and ssp. over the world. Possibly derived from var. nipposinca form the E Asiatic branch ramosa DC. in the area of Brussels and (Diederichsen 2001). Toxopeus and Oost distributed from there since about 1785 (1985) proposed a grouping into cultivar (Helm 1959). We assume an involvement of groups. convar. acephala and var. sabauda (Gladis ssp. sylvestris (Lam.) Janchen, Janchen et and Hammer 2003). Wendelbg. Wild progenitor, today rare and Brassica rapa L. em. Metzg. difficult to find. The common weedy races In the case of B. rapa (the Brassica also belong here (var. sylvestris (Lam.) campestris cytodeme according to Harberd Briggs). 1972), primary domestications may have f. campestris (L.) Bogenhard. Wild and happened in W Asia/ E Mediterranean weedy forms, widely distributed but where the wild progenitor ( B. rapa ssp. becoming rare with the reduction of oilseed

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 460

turnip cultivation. Many transitions to the for which provisional names have been following subspecies. proposed for better comparison and to fit ssp. oleifera (DC.) Metzg. Oilseed turnip. into the general treatment of Brassica rapa. var. oleifera DC. Oilseed crop. convar. rapa (incl. convar. rossica (Sinsk.) f. praecox (DC.) Mansfeld. Spring oilseed Scheb., convar. europaea Scheb.). Variable turnip. European group. Today often a relic crop. f. autumnalis (DC.) Mansfeld. Winter var. rapa. With grey and black tubers. A oilseed turnip. typical relic crop in Central Europe is e.g. var. ruvo (Bailey) comb. prov.(Brassica ruvo the ‘Bayerische Rübe’(Reiner et al. 2005). Bailey; Yang and Quiros (2010) see var. teltoviensis Alef. Another relic crop in relations to ssp. nipposinica). This is a western and central Europe with small common vegetable e.g. in S Italy (cime di tubers, e.g. ‘Teltower Rübchen’ (Landsrath rapa) and in Azerbaijan or Turkey (turp). and Hammer 2007). The young sprouts with the buds are var. septiceps Bailey. Seven-top turnip or eaten as a vegetable. There is rich variation Italian kale. The well developed are in the area with many different types. Used usually not eaten but the young sprouts, in the same way as broccoli or cauliflower similar to var. ruvo. This turnip comes from (Astley et al. 1984, Maly et al. 1987, Gladis Italy and has a restricted growing area in the and Hammer 1992, Hammer et al. 1999). It , Canada and . Placed is an interesting case for the evolution of here in spite of tubers not used (Šebalina and inflorescenses as vegetable and should be Sazonova 1985). classified as a botanical variety var. rossica Scheb. Relic crop in parts of independently of var. oleifera. Eastern to Western Europe. ssp. rapa (incl. ssp. orientalis (Sinsk.) var. rubra (Sinsk.) Scheb. Red tubers, in Scheb., ssp. mesopotamica Scheb., ssp. eastern and northern parts of Europe. afghanica (Sinsk.) Scheb., ssp. japonica var. europaea Scheb. Turnip with white Scheb.). Turnip. Widely grown as a root tubers, still wide-spread in Europe. vegetable. In E Asia also the leaves are var. flava (Sinsk.) Scheb. Turnip with eaten, this has been also observed in N yellow tubers, wide-spread in Europe and Africa (Hammer and Perrino 1985). After beyond (e.g. ). Hanelt (1986) the most variable race of convar. orientalis (Sinsk.) comb. prov. (B. Brassica rapa. From Japan also races with campestris var. orientalis Sinsk.). Group laciniated leaves are reported (Kitamura with small tubers from Turkey and the 1950, Shebalina and Sazonova 1985). In Middle East (Šebalina and Sazonova 1985). China they are known since the first Century var. orientalis (Sinsk.) comb. prov. Only one of our era (Diederichsen 2002). Alefeld botanical variety. (1866) described the races present in convar. mesopotamica (Scheb.) comb. prov. European gardens of his time and mentioned (B. rapa ssp. mesopotamica Scheb.), Group material from E Asia. Many types are from Irak. already lost in Europe. The same may be var. mesopotamica Scheb. Turnips with true for material from a postulated Central green head. Asian center of diversity (Sinskaja 1969). var. violascens Scheb. Turnips outside Šebalina (1968), considering the turnip as a violet, inside white. species, classified it into five subspecies and convar. afghanica (Sinsk.) Scheb., incl. many further infraspecific entities. They are convar. chinensis Scheb., convar. indica taken as the basis for the present treatment (Sinsk.) Scheb., convar. ferganica (Sinsk.)

461 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Scheb., convar. sinensis (Sinsk.) Scheb. Helm (1963 b) summarizes the differential Variable group from Afghanistan, N India, characters from the first variety, which are Uzbekistan, Kirgizstan and W China, only in not considered as sufficient (Gladis and rare cases known outside this area. Hammer 1992). But they can be taken as a var. afghanica (Sinsk.) Scheb. From working basis. Afghanistan and Uzbekistan. var. rosularis (Tsen et Lee em. Lee) Hanelt. var. indica (Sinsk.) Scheb. The characteristic Already Lee (1982) included B. narinosa seed color is yellow brown, going to be here. For further discussions see Sun (1946) diplaced by varieties from the West and Lin (1980). (Šebalina and Sazonova 1985). var. dubiosa (Bailey) comb. prov. var. hybrida Scheb. Variety with variable Described by Bailey (1922, 1930) from characters. China (district of Nanjing). Similar to var. var. ferganica Scheb. Turnips from rosularis and also to var. albiflora (Gladis Uzbekistan and Kirgizstan. and Hammer 1992). var. sinensis (Sinsk.) comb. prov. (B. ssp. pekinensis (Lour.) Hanelt. Pe-tsai, campestris var. sinensis Sinsk.). Turnips . Cultivated in China since from W China. more than 500 years (Bretschneider 1898). convar. japonica (Scheb.) comb. prov. (B. A head-forming race with a tendency for rapa ssp. japonica Scheb., B. rapoasiatica wider distribution in the last 100 years ssp. japonica Sinsk., nom. illeg., B. rapa ssp. (Hanelt 1986a). japonica (Sinsk.) Scheb., comb. inval.). var. pandurata (Sun) Gladis. Fruit- and Group from Japan and E China. seed-characters show a relation to Indian var. japonica Scheb. Japan, also known in material (Gladis and Hammer 1992). the Western World (Šebalina and Sazonova var. laxa (Tsen et Lee) Hanelt. With loose 1985). heads. var. alborosea Scheb. var. glabra E. Regel (B. chinensis var. var. intermedia Scheb. pekinensis (Lour.) Sun). There are different ssp. chinensis (L.) Hanelt (incl. ssp. narinosa head forms which are formally described as (Bailey) Hanelt) pak-choi, Chinese mustard. varieties of B. pekinensis (Helm 1961). Sun There have been several attempts to describe (1946) as well as Gladis and Hammer the variation of this subspecies (Tsen and (1992) recommend no further splitting. It Lee 1942, Helm 1963 b, Lee 1982, Gladis might be done on the level of forms within and Hammer 1992). However, there is still var. glabra as follows: no generally excepted infra-subspecific f. laxa (Tsen et Lee) comb. prov. with system. Typical crop from E Asia. upright, loose heads Important races are: f. pekinensis (Helm) comb. prov., with var. chinensis (Juslen.) Kitam. This race is compact round or elongated heads, and the most typical one. f.. cylindrica (Tsen et Lee) comb. prov. This var. communis Tsen et Lee. For tradional highly domesticated race, which has been conservation the leaves are dried or salted. developed from var. glabra, experienced a Occasionally tuberous roots are formed. wide distribution in Europe and America. There are reports about the consumption of ssp. nipposinica (Bailey) Hanelt. A young sprout. There are other variations vegetable crop from E. Asia with rather which are in need of additional studies. variable leaf characters (simple and laciniate var. parachinensis (Bailey) comb. prov. (B. leaves). chinensis var. parachinensis (Bailey) Sinsk.)

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 462

var. perviridis Bailey. (ssp. perviridis (Ethiopia). The B genome turned out to be Bailey). Tendergreen, mustard. relatively conservative (Liu and Wang Cultivated as a and also for 2006). Brassica nigra is an ancient crop the tuberous roots. Introduced in the first plant with interesting resistance characters half of the last century to SE United States. (Sheng et al. 2012). Sinskaja (1928) Related to B. chinensis var. utilis Tsen et differentiates between a Western and a Lee (1942), see under ssp. trilocularis. Eastern group, which can be accepted as var. lorifolia Bailey. The tuberous roots are subspecies. In Italy formerly frequently used. Possibly related to var. chinoleifera cultivated (Maly et al. 1987), now mostly a (Gladis and Hammer 1992). weed (Aeschimann et al. 2004). A third var. dissecta (Schulz) Gladis. All races with group from India (var. indica Sinsk.) is not dissected leaves (Makino 1912) belonging to yet sufficiently known (Gladis and Hammer ssp. nipposinica are placed here. 1992). ssp. trilocularis (Roxb.) Hanelt. Oilseed ssp. nigra. Western group. Cultivated in crop, only occasionally vegetable. Mainly Europe, Ethiopia, Afghanistan, Crete and used in India. Cyprus. var. trilocularis (Roxb.) Duthie et Fuller. var. nigra (incl. var. torulosa (Pers.) Alef., Variable oilseed crop, showing some var. turgida (Pers.) Alef.). Also common crossing barriers to other subspecies of the ruderal and segetal plant. species (Olsson 1954). Yellow sarson is var. pseudocampestris Sinsk. Segetal plant self-compatible. Very old oilseed crop, adapted to Brassica rapa ssp. oleifera. already mentioned in Sanskrit documents Today very rare because of the decline of B. from 1500 BC. (Diederichsen 2001). Fruit rapa ssp. oleifera cultivation. with 3 and 2 valves. var. abyssinica A. Braun. Cultivated and var. quadrivalvis (Roxb.) Duthie et Fuller. weedy in Ethiopia. The only known cruciferous crop with 4- ssp. hispida (Schulz) Gladis. Eastern group. valved fruit, sometimes containing a second Cultivated in , Israel, Thessalia and small fruit within the basal siliques. The Asia Minor. vernacular name yellow sarson is also var. rigida Sinsk. High growing plants from applied to this variety. Syria, Israel and Thessalia. var. chinoleifera (Viehoever) Kitam. (var. var. orientalis Sinsk. Low growing plants utilis Tsen et Lee). Variable oilseed crop from Asia Minor. from China, especially in leaf- and seed- Brassica tournefortii Gouan colour (Tsen et Lee 1942). By Gladis and Cultivated on a small scale in NW India and Hammer (1992) placed under ssp. W Tibet as an oilcrop. The chromosome nipposinica. But clearly similar to the Indian number is n = 10, but cytogentically ssp. trilocularis. different from Brassica rapa (n = 10). ssp. dichotoma (Roxb.) Hanelt. Brown Brassica carinata A. Braun sarson, toria. The most traditional oilseed Abyssian mustard is grown as a oilseed crop crop of India. Toria is close to ssp. oleifera and also as a leaf vegetable in Ethiopia, (Olsson 1954). Disappearing or already Kenia and some other E African countries disappeared from cultivation. (Alemayehu and Becker 2002). Today also Brassica nigra (L.) Koch as a new crop in Europe and America Black mustard is mostly cultivated as an (Warwick et al. 2006). The variation of this oilseed crop, for the production of mustard, crop is not very high. As the two parental occasionally grown as a vegetable species B. oleracea and B. nigra possibly

463 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

have been introduced to Ethiopia with ssp. napus. Mostly oilsed crop and fodder Amharic agricultural tribes in the first plant, one race is grown as a vegetable. First millennium BC, B. carinata could have its reliable reports in herbals from the 16th and origin here and was later distributed to other 17th centuries (Diederichsen 2001). Possibly African areas. In Italy in trials as an originated in the W Mediterranean area. alternative oil crop for biodiesel production Wild (weedy?) plants have been reported (Cardone et al. 2003). from N Africa (Fiori 1923/25). Brassica napus L. This allopolyploid hybrid var. napus. Rape, canola. Very common as (x = 19) with genomes from Brassica rapa an oilseed crop in many parts of the world, and B. oleracea (Hu et al. 2011) evidently recently also in E Asia (see Diederichsen originated under domestication. As place of and McVetty 2011). origin N Africa has been proposed, were f. annua (Schübl. et Mart.) Thell. Summer “truly wild” material was found (Fiori 1923- rape. 25). Brassica napus is considered as a f. biennis (Schübl. et Mart.) Thell. Winter relatively young crop that originated in rape. limited geographical region (Kimber and var. pabularia (DC.) Reichenb. Leaf rape, McGregor 1995). There are differences in Siberian kale. A variable leaf vegetable. the genetic background between Eastern and In addition to the described diversity, there European genotypes (Zhou et al. 2006). have been done several artificial Natural crosses between the two parental recombinations of this amphidiploid from species have been observed (Hansen et al. different (con-)varieties of the progenitor 2001). Subspecies napus is mainly grown as species, e.g. an oilseed crop and became a world crop in B.-×-napus-cauliflower and broccoli the last century (Diederichsen and McVetty developed from B. oleracea convar. botrytis 2011). Subsp. rapifera is mainly a root + B. rapa var. ruvo, and vegetable. B.-×-napus-cabbage developed from B. ssp. rapifera Metzg.(ssp. napobrassica (L.) oleracea var. capitata + B. rapa var. glabra Hanelt). Swede, Swedish turnip, . (namend Hakuran) and others (see Clauss Formerly concentrated on Europe, especially 1975, Yamagishi and Takayanagi 1982).. N Europe and , today also in many Today, hakuran is available in the market as other areas as the United States, Siberia and vegetable crop. The formal classification of Far East. A separate origin from ssp. napus the resynthesis rape vegetables is missing has been proposed (Diederichsen 2001). In yet. the classification of swede we follow Brassica juncea (L.) Czern. This alloploid Šebalina and Sazonova (1985). species derived from Brassica rapa and B. var. rapifera Metzg. European races. Alefeld nigra. Archaelogical remains from 2.000 BC (1866) described a number of varieties. The have been found in the Indus valley N European history has been stressed by (Diederichsen 2001). The Middle East is Ahokas (2002). A possible origin is in the most likely the area of origin, where both of Mediterranean area, where both progenitors the original species occur. The largest co-existed (Hanelt 1986 a). Many cultivars. variation of B. juncea today is found in W var. rossica Sinsk. Russia to . and Central China (Huangfu et al. 2009) and var. sibirica (Scheb.) Bondar. et Pivovar. Central Asia (Rabbani et al. 1998). Yellow swede from Siberia. Informally they are called root mustard, var. schebalinae Bondar. et Pivovar. White stem mustard, leafy mustard, stalk mustard swede from Siberia. and seed mustard (Fu et al. 2006).

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 464

Vegetable types originated earlier than oil ssp. integrifolia (West) Thell. A common types, but there is no conclusion concerning leaf vegetable in E and SE Asia. In Italy the development of oil seed types from also cultivated as an oil seed (Fu et al. vegetable types (Wu et al. 2009). The 2006), in rare cases as a leaf vegetable infraspecific variation has been classified by (Branca 2004). Bailey 1922, Sinskaja 1928, Sabnis and var. subintegrifolia Sinsk. In this group the Phatak 1935, Tsen and Lee 1942, Helm use of seeds is predominant. Sinskaja (1928) 1959, Nishi in Tsunoda et al. 1980, Chen names the following varieties: 1982. We follow here mainly the treatment var. sareptana Sinsk. of Gladis and Hammer (1992), which has var. subintegrifolia Sinsk. been also accepted by Diederichsen (2001). var. subsareptana Sinsk. Brassica juncea includes vegetables and var. subcrispifolia Sinsk. oilseed crops. Formerly Brassica cernua var. mongolica Sinsk. (Poir.) Fourb. et Hemsl. was considered as a var. japonica Bailey separate species containing the oilseed var. multisecta Bailey brassicas. var. integrifolia (Stokes) Sinsk. Basal leaves ssp. napiformis (Paill. et Bois.) Gladis. The not dissected. Collected by us in Korea root tubers are used as vegetable. Several (Gladis and Hammer 1992). races have been described (var. megarrhiza var. rugosa Roxb. ex Prain (incl. var. Tsen et Lee (Fu et al. 2006), var. napiformis agrestis Prain, var. typica Prain, var. (Paill. et Bois.) Kitam., var. multisecta cuneifolia (Roxb.) Prain). Cultivated as leaf Baranov), but despite of additional material vegetable and oil seed. from Korea, which could be studied (Kim et var. capitata Tsen et Lee ex Lee. Races al. 1987), the high variation found in still forming heads are already described by limited material does not allow for deeper Kumazawa and Abe (1955/56). insights. Interesting figures of several types var. longidens Bailey (var. strumata Tsen et have been provided by Qi et al. (2007). This Lee). Leaves appear unsymmetrical. group is urgently in need for a revision. var. crispifolia Bailey. Vegetable with curly ssp. tsatsai (Mao) Gladis. Group with fleshy leaves, sometimes ornamental. stalks used as vegetable mainly from var. celerifolia Tsen et Lee. Doubtfully Sichuan. belongs to this species (Gladis and Hammer var. multiceps Tsen et Lee. Many fleshy 1992, see also Yang-Zheng 1986). short stalks. Possibly related to var. ssp. juncea. Mainly cultivated for the seeds, gemmifera Lin et Lee (Lin and Lee 1985). sometimes as fodder plant. var. urophylla Tsen et Lee. Described from var. juncea. Important oilsed plant, China (Tsen and Lee 1942) and similar to occasionally grown as a leaf vegetable (Fu var. multiceps. Collected by Hammer 1986 et al. 2006) and ornamental plant. in Cuba as a ruderal plant, in an area, where var. brachycapa Thell. (var. lonigpes Tsen et Chinese immigrants played a role as Lee, var. orthocarpa Sun). Oilseed plant and horticulturalists (Hammer et al. 1992 – also fodder crop. 1994). Exact determination not yet possible: var. linearifolia Sun. Plants with reduced var. chirimenna (Makino) Burkill. Oilseed leaf blade (Sun 1946, Lee 1982). plant. var. tsatsai Mao(var. tumida Tsen et Lee). var. edona Makino. A of var. Variable vegetable in Szechuan where more rugosa? than 11 forms or cultivars occur (Lee 1982).

465 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

var. facilifolia Li. The young inflorescenses another ecological possibility for surviving are used as vegetable. on calcareous cliffs close to the side, Brassica ×harmsiana O.E. Schulz mainly in the Mediterranean area, where it Synthetic alloploid hybrids with one C could escape goats and fire (Perrino et al. genome of Brassica oleracea and at least two 1992). But also these wild plants can be A genomes of Brassica rapa (Diederichsen derived from cultivated ones as seems to be 2001). A number of similar synthetic the case with the Atlantic populations of material have been produced for new oil and Brassica oleracea (Maggioni et al. 2010). All fodder plants, sometimes for vegetable crops the other cultivated species of Brassica (Frandsen and Winge 1932, Gladis and chose a different pathway. They escaped in Hammer 1990). Analogous hybrids with the agricultural areas and became segetal one genome of Brassica rapa and at least and ruderal plants. The large fields of two genomes of Brassica oleracea are cultivated plants (mostly for oil crops) but introduced under the name of Brassica also garden plants as good pollen donors ×napoleracea Chiang in Cruciferae (mostly for vegetable races) provided a great Newsletter 10 (1985) 25. genetic influence on the weedy or ruderal Comprehensive keys for botanical progenitors, changing their genetic determinations of the cultivated species and constitution. The results are populations their infraspecific items have been provided closely related to the cultivated races, which by Bailey (1949) and Gladis and Hammer are different from the wild progenitor of (2003). Gladis and Hammer (1992) also crops. This behavior of the cultivated races considered the wild and weedy races. Other is called “genetic aggression” (Hammer keys are confined to species (e.g. Šebalina 2003). Ruderal and weedy races of nearly all and Sazonova 1985). cultivated Brassica species are widely distributed in the growing areas and can WILD AND WEEDY PROGENITORS serve as useful genetic resources (see last AND DESCENDENTS paragraph). Spontaneous outcrossings resulting in genetic transfer (introgessions) from the LANDRACES AND FURTHER cultivated to the wild species and vice versa EVOLUTIONL have been mentioned by many authors (see There is a close relationship between the compilation by Warwick et al. 2009). Such evolution of crops plants and human cultural spontaneous gene transfer has received evolution (Maffi 2001). We still lack a broad increased attention after the introduction of approach of this comprehensive interrelation transgenic cultivars of , B. napus despite the great achievements reached by var. napus, during the 1990s (Beckie et al. A. De Candolle (1806 – 1893), Ch. Darwin 2006) and included many other crops. (1809 – 1882) and N.I. Vavilov (1887 – The wild progenitor and the beginnings of 1843). At the beginning of scientific plant domestication are often unclear and still breeding agri- and horticulture were full of under discussion (see remarks under the morphologically distinguishable landraces. different species), because of the lack of Their value for ongoing was archaeological material and the difficulties appreciated at a relative early time of its identification. Linguistic and (Proskowetz and Schindler 1890). The ethnological data can often provide some gradual displacement of these landraces by first indications (Zohary et al. 2012). In breeding varieties soon ewaked concern of Brassica oleracea the wild progenitor found farsighted scientists, but only relatively late

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 466

the term “genetic erosion” was coined breeders and producers, completely (Bennett 1968). Soon the “plant genetic neglecting the landraces (Pickersgill and resources movement” developed (Pistorius Karamura 1999). We cannot go into details 1997). More and more landraces were here, but we want to stress the usefulness of disappearing despite of great efforts to infraspecific formal classifications for crop maintain them ex situ and, in the last twenty plant taxonomy, evolution and history. years, on farm. Morphological classification Table 4 shows the high number of of cultivated plants was the primary infraspecific races in some of the Brassica- approach from the time of Linnaeus (1753, species. Some species are characterized by 1763). He was aware of variation under low numbers of infraspecific races, as human influence and placed cultivated Brassica carinata. In other cases the low race plants together with the wild ones in species, number may be due to unsufficient indicating them by Greek letters. This knowledge (e.g. Brassica juncea in E Asia). method was maintained for a long time, e.g. At any rate, infraspecific classifications by A.P. de Candolle (1822) and Metzger allow an overview on the available material (1833). Alefeld (1866) advocated a similar and their variation in the fields and in treatment of wild and cultivated plants. His collections. They are useful for historic only concern was the large variation of studies and provide a good method for cultivated plants. He tried to solve this showing gaps in material and areas. Their problem by the introduction of a new special usefulness can be seen for the infraspecific category (later proposed as characterization of landraces. convar.) for groups of botanical varieties. There is a general trend in landraces towards This system was further elaborated and their flourishing in the 19th/20th centuries turned out to be applicable for landraces. In and their fading away at an increasing speed Brassica this system was used by Bailey from the second half of the 20th century. from the United States (1922 – 1942), the The speed of this process is high, but even in Vavilov school of Russia (e.g. Sinskaja developed countries niches can be found in 1928, Šebalina and Sazonova 1987), the which an evolution of landraces proceeds. Mansfeld school of Gatersleben (Germany) The domestication syndrome (Hammer (e.g. Helm 1959 – 1964). The system is still 1984, 2003, Pickersgill 2009) is further used in historical context, to classify diverse developing. The nature of selection during genbank collections, to report about plant domestication changed (Purugganam evolutionary trends in cultivated plants, and Fuller 2009). Highly domesticated including genetic erosion (e.g. Hammer and crops, obtained by special plant breeding Laghetti 2005) and for other research on methods, resulted in high yielding varieties genetic resources. For plant breeding and (Gepts 2004, Brown 2010). The concept of plant production it is less adapted, mainly super-domestication shows the way from because of the tremendous loss in plant breeding to crop engineering (Vaughan morphological variation during conscious et al. 2007). Super-domestication and plant breeding. Several times a schism was gigantism can be also observed in Brassica. proposed, to separate the classifications of Giant cabbages have been decribed from wild and cultivated plants, one of the latest Turkey (Zhukovsky 1933) and hybrids of efforts is the creation of “cultonomy” vs. Brassica rapa ssp. pekinensis are highly “taxonomy” (Hetterscheid and Brandenburg productive. The amphidiploid Brassica spp. 1995). Accordingly the ICNCP (Brickell et as the world crop Canola (Brassica napus) al. 2009) was adapted mainly for the use of and many of the newly generated

467 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

amphidipoids of the Brassica ×harmsiana primary products perform agronomically type show a fixed heterosis and we begin to very poorly and have to be improved by understand the mechanisms of gene prebreeding methods (Seyis et al. 2003). expressions (Osborn et al. 2003). But the

Table 4. Cultivated brassicas and their infraspecific items according to formal classification Taxa ssp. convars bot. varieties other races Brassica oleracea ssp. oleracea 1 ssp. bourgaei 5 ssp. cretica 4 ssp. hilarionis 1 ssp. incana 4 ssp. insularis 6 ssp. macrocarpa 2 ssp. robertiana 1 ssp. rupestris 3 ssp. villosa 4 ssp. capitatoides 1 ssp. capitata 6 18 15 Brassica rapa ssp. sylvestris 1 ssp. oleifera 2 2 ssp. chinensis 5 ssp. pekinensis 4 ssp. nipposinica 3 ssp. trilocularis 2 ssp. dichotoma 1 Brassica nigra ssp. nigra 3 ssp. hispida 2 Brassica tournefortii 1 Brassica carinata ssp. rapifera 4 ssp. napus 2 2 Brassica juncea ssp. napiformis 3 ssp. tsatsai 4 11 ssp. subintegrifolia 14 ssp. juncea 2

GENETIC RESOURCES Genetic exchange is possible among several The still available rich material of landraces, cultivated and wild taxa of the genus wild and weedy races is seen as the basis for Brassica despite different chromosome further crop evolution in Brassica (table 5,). numbers. This became recently very According to the gene pool concept of important with the possibility of unintended Harlan and De Wet (1971) there are three introgressions of GMOs (Beckie et al. genepools. Genepool three is very much 2006). Species that have the same enlarging at present, so that Hammer (1998) chromosome numbers and readily cross with and Gepts and Papa (2003) proposed a each other, form a cytodeme (Harberd 1972) fourth genepool for the new products of or crossing group (first genepool). The term plant transformation and genomics. A new Brassica coenospecies (more or less second concept considers an enlarged third genepool) encompasses all species that are genepool. The fourth genepool should capable of exchanging genes with the contain any synthetic strain shown with nucleic acid frequencies, DNA and RNA, that do not occur in nature (Gladis and Hammer 2002).

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 468

Table 5. Summary of important cultivated species of the genus Brassica L., their infraspecific diversity and selected common names Species Karyotype1 Subspecies Further infraspecific grouping Usage type Common name B. rapa L. em. Metzg.(syn. B. n=10, A ssp. oleifera (DC.) Metzg. Seed oil, forage, green manure Oilseed turnip, turnip rape, field campestris L.) mustard, canola n=10, A ssp. oleifera (DC.) Metzg. Leaf and inflorescence as vegetable Cime di rapa (Italian) n=10, A ssp. trilocularis (Roxb.) Hanelt Seed oil, sometimes vegetable Yellow sarson n=10, A ssp. dichotoma (Roxb.) Hanelt Seed oil, vegetable, forage Brown sarson, toria (Hindi), Indian rape n=10, A ssp. rapa Root vegetable, forage Turnip n=10, A ssp. chinensis (L.) Hanelt Leaves and petioles as vegetable Pak-choi n=10, A ssp. pekinensis (Lour.) Hanelt Leaves and leafy rosettes as vegetable Pe-tsai, Chinese cabbage n=10, A ssp. nipposinica (Bailey) Hanelt Leaves as vegetable Mizuna (Japanese) B. nigra (L.) Koch n=8, B ssp. nigra Seed oil and condiment, vegetable and Black mustard (western type) forage n=8, B ssp. hispida (Schulz) Gladis Seed oil and condiment, vegetable and Black mustard (eastern type) forage B. oleracea L. n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. gongylodes L. Stem as vegetable Kohlrabi n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. medullosa Thell. Stem as forage Marrow-stem kale n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. palmifolia DC. Leaves as vegetable Palm kale n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. sabellica L. Curled leaves as vegetable Curled kitchen kale n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. selenisia L. Leaves as vegetable Russian kale n=9, C ssp. capitata (L.) DC. convar. acephala (DC.) Alef. var. viridis L. Leaves as vegetable or forage Colewort, collard n=9, C ssp. capitata (L.) DC. convar. botrytis (L.) Alef. var. botrytis L. Budding inflorescence as vegetable Cauliflower n=9, C ssp. capitata (L.) DC. convar. botrytis (L.) Alef. var. italica Plenck Budding inflorescence as vegetable Broccoli n=9, C ssp. capitata (L.) DC. convar. botrytis (L.) Alef. var. alboglabra (Bail.) Sun Budding inflorescence as vegetable Chinese kale n=9, C ssp. capitata (L.) DC. convar. capitata (L.) Alef. var. capitata L. Leafy heads as vegetable Cabbage n=9, C ssp. capitata (L.) DC. convar. capitata (L.) var. sabauda L. Leafy heads as vegetable Savoy cabbage n=9, C ssp. capitata (L.) DC. convar. costata (DC.) Gladis var. costata DC. Petioles and leaves as vegetable Tronchuda kale n=9, C ssp. capitata (L.) DC. convar. costata (DC.) Gladis var. helmii Gladis et Hammer Ornametal leaf type Stor ribbekaal (Danish) n=9, C ssp. capitata (L.) DC. convar. fruticosa (Metzg.) Alef. var. ramosa DC. Leave vegetable and forage Branched, vegetative kale n=9, C ssp. capitata (L.) DC. convar. gemmifera (DC.) Gladis var. gemmifera DC. Young side buds as vegetable Brussel's sprouts n=9, C ssp. capitata (L.) DC. convar. gemmifera (DC.) Gladis var. polycephala Thell. Leafy, headed side branches as Branched cabbage vegetable B. juncea (L.) Czern. n=18, AB ssp. juncea Seed oil, forage Indian mustard, oriental mustard, rai (Hindi) n=18, AB ssp. integrifolia (West) Thell. Leaf vegetable Leaf mustard n=18, AB ssp. napiformis (Paill. et Bois) Root vegetable Tuberous-rooted mustard Gladis n=18, AB ssp. tsatsai Mao Petioles and leaves as vegetable Tsatsai B. napus L. n=19, AC ssp. napus var. napus Seed oil, forage, green manure Rapeseed, canola n=19, AC ssp. napus var. pabularia (DC.) Leaf vegetable Nabicol, rape-kale Rchb. n=19, AC ssp. rapifera Metzg. (ssp. Root vegetable and forage Swede, rutabaga napobrassica (L.) Hanelt) B. carinata A. Braun n=17, BC Seed oil and vegetable use Abyssinian mustard 1 Karyotype according to Mizushima (1980).

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 469

important cultivated Brassica species the 18th Crucifer Workshop “Exploitation of (Prakash et al. 1999). Brassica Diversity for Improving A wide range of species can be employed Agriculture Chains”, Catania 12 – 16 for crop improvement (third genepool). November 2012.We thank Prof. Dr. F. Prebreeding or germplasm enhancement is Branca for his support. employing an always increasing tertiary genepool. But in the breeding programmes REFERENCES the tendency is to stay within the genepool Gornall R.J. 1983. Recombination systems of adapted material (Downey and Rakow and plant domestication. Biol. J. Linn. 1987, Becker et al. 1999). Soc. 20, 375 – 383. The pollination mode of the different Aeschimann D., Lauber K., Moser D.M. and Brassica species ranges from self Theurillata J.-P. 2004. Flora alpina (Ein incompatibility to complete self-fertility. Atlas sämtlicher 4500 Gefäßpflanzen der Even within species there exist variations Alpen). 3 vols. Haupt Verlag, Bern, among plant types for pollination mode Stuttgart, Wien. (Gladis and Hammer 1992). As a tendency, Ahokas H. 2002. Cultivation of Brassica the diploid species are obligate outcrossing, species and by ancient Finnic while the amphidiploid bastards are self people, traced by linguistic, historical fertile (Downey and Rakow 1987). and ethnobotanical data; revision of Large genebank collections for Brassica Brassica napus as B. radice-rapi. Acta germplasm have been assembled in Europe, Bot. Fenn. 172, 1 – 32. India and China. For wild species of the Alefeld F. 1866. Landwirthschaftliche Flora. genus Brassica an important collection is Wiegandt & Hempel, Berlin, 363 pp. available in Spain (Gómez-Campo 2005). Alemayehu N. and Becker H.C. 2002. The World Vegetable Center (AVRDC) in Genotypic diversity and pattern of Taiwan maintains Brassica with 1390 variation in a germplasm material of Brassica accessions (AVRDC 2009). Ethiopian Mustard (Brassica carinata A. Gladis and Hammer (1990, 2003) provided Braun). Genet. Resour. Crop Evol. 49, information on techniques for germplasm 573 – 582. regeneration and preservation. Ambrosoli M. 1997. The wild and the sown. The enlarged tertiary genepool includes the Botany and agriculture in Western related genera , Diplotaxis, Eruca, Europe: 1350 – 1850. Cambridge Erucastrum, , Sinapis, University Press, Cambridge. Sinapidendron, Trachystoma, Andersen N.S., Poulsen G., Andersen B.E., Enarthrocarpus, Raphanus, , Kiaer L.P., D’Hertefeldt, Wilkinson Pseuderucaria and Rytidocarpus and M.J.and Jørgensen R.B. 2009. Processes successful introgressions have been reported affecting genetic structure and (Hu et al. 2009). As already stated in the conservation: a case study of wild and introduction of this paper, a first recorded cultivated Brassica rapa. Genet. Resour. successful wide cross resulted in Raphanus Crop Evol. 56, 189 – 200. sativus × B. oleracea (Sageret 1986). Andrews S., Leslie A. and Alexander C. (eds) 1999. Taxonomy of Cultivated ACKNOWLEDGEMENTS Plants: Third International Symposium. This is a slightly elaborated version of a Proc. Meet. Edinb. 1998. Royal Bot. keynote paper presented at the 6th Gard., Kew. International Symposium of Brassica and

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 470

Argentieri M.P., Accogli R., Fanizzi F.P. Exploration, Utilization and and Avato P. 2010. Glucosinolates Conservation of Plant Genetic profile of “Mugnolo”, a variety of Resources. FAO, Rome. Brassica oleracea L. native to southern Beridze R.K., Hanelt P. and Fritsch R. 1983. Italy (Salento). Planta Medic. , 494 – Report of a collecting mission to the 999. Georgian SSR 1982 for the study of Astley D., Crisp P. and Perrino P. 1984. indigenous material of cultivated plants. Cruciferous crops in Italy. The collection Kulturpflanze 31, 173 – 184. of landraces of cruciferous crops in Bothmer R. von, Gustafsson M. and EC Countries. Final Rep. EC Res. Snogerup S. 1995. Brassica sect. Progr.0890, IVT. Brassica (Brassicaceae). II. Inter- and AVRDC 2009. The World Vegetable intraspecific crosses with cultivars of Center. http://www.avrdc.org/index.html Brassica oleracea. Genet. Resour. Crop Bailey L.H. 1922. The cultivated Brassicas Evol. 42, 165 – 178. I. Gentes Herbarum 1, fasc. 2, 51 – 108. Boukema I.W. and van Hintum T.J.L. 1999. Bailey L.H. 1930. The cultivated Brassicas Genetic Resources. In: Gómez-Campo, II. Gentes Herbarum 2, fasc. 5, 211 – C. (ed.) Developments in plant genetics 267. and Breeding, 4. Biology of Brassica Bailey L.H. 1940. Certain noteworthy coenospecies. Elsevier, Amsterdam, pp. Brassicas. Gentes Herbarum 4, fasc. 9, 461 - 479. 322. Branca F. 2004. Esperienze sull’impiego di Bailey L.H. 1949. Manual of cultivated Brassica macrocarpa per il controllo dei plants most commonly grown in the nematode galligeni nel pomodoro. Atti continental United States and Canada, Workshop Internazionale “Produzione in rev. ed. Macmillan, New York. serra dopo l’era del bomuro di metile”, Bailey L.H. and Bailey E.Z. 1976. Hortus tenuto a Comiso nell’aprile 2004. Third. New York, London, 1290 pp. Branca F. and Iapichino G. 1997. Some wild Baillargeon G. 1986. Eine taxonomische and cultivated Brassicaceae exploited in Revision der Gattung Sinapis Sicily as vegetables. FAO/IBPGR Plant (Cruciferae: Brassicaceae). 268 pp., Genet. Res. Newsl. 110, 22 – 28. Berlin. Branca F., Ragusa L., Argento S. and Becker H., Löptin H.and Röbbelen G, 1999. Tribulato A. 2010. Attivitá per la Breeding: An Overview. In: Gómez- conservazione in situ di specie spontanee Campo, C. (ed.) Developments in plant del genero Brassica (n=9) in Sicilia. IV genetics and breeding, 4. Biology of Convegno Nazionale Piante Brassica coenospecies. Elsevier, Mediterranee “Le potenzialità del Amsterdam, pp. 413 - 460. territorio e dell’ambiente”, pp. 175 – Beckie H.J., Harker K.N., Hall L.M., 180. Warwick S.I., Légère A., Sikkema P.H., Bretschneider E. 1898. History of the Clayton G.W., Thomas A.G., Leeson European Botanical Discoveries in J.Y., Séguin-Swarts and Simard M.-J. China I, II. Leipzig. 2006. A decade of herbicide-resistant Brickell C.D., Alexander C., David J.C., crops in Canada. Can. J. Plant Sci. 86, Hetterscheid W.L.A., Leslie A.C. 1243 – 1264. Malecot V. and Xiaobai J. 2009. Bennett E. 1968. Record of the FAO/IBP International Code of Nomenclature for Technical Conference on the

471 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Cultivated Plants. ISHS Scripta Crisp P. and Gray A.R. 1984. Breeding old Horticulturae 10, 204 pp. and new forms of purple heading Brown A.H.D. 2010. Variation under broccoli. Cruciferae Newsletter 9, 17 – domestication in plants: 1859 and today. 18. Philos. Trans. R. Soc. Lond. B, Biol. Sci. Daun J. K. Eskin N.A.M. and Hicking D. 365, 2523 – 2530. (eds), 2011. Canola: Chemistry, Cardone M., Mazzoncini M., Menini S., Production, Processing and Utilization. Rocco V., Senatore A., Seggiani M. and AOCS Press, Urbana. Vitolo S. 2003. Brassica carinata as an De Candolle Alphonse 1883. L’origine des alternative oil crop for the production of plantes cultivées. Paris. biodiesel in Italy: agronomic evaluation, De Candolle Augustin Pyramus 1824. fuel production by transestrification and Memoir on the different species, races characterization. Biomass and Bioenergy and varieties of the genus Brassica 25, 623 – 636. (cabbage) and the genera allied to it, Chen H.G. and Wu J.S. 2008. which are cultivated in Europe. Transact. Characterization of fertile amphidiploids Hort. Soc. London 5, 1 – 43 (orig. in between Raphanus sativus and Brassica French 1822, in German 1824). alboglabra and the crossability with Dias J.S. 1995. The Portuguese tronchuda Brassica species. Genet. Resour. Crop cabbage and galega landraces: A Evol. 55, 143 – 150. historical review. Genet. Resour. Crop Chen S.-R. 1982. The origin and Evol. 42, 179 – 194. differentiation of mustard varieties in Dias J.S. 2012. Portuguese kale: a relic leafy China. Cruciferae Newsl. 7, 7 – 10. vegetable crop. Genet. Resour. Crop Christensen S., von Bothmer R., Poulsen G. Evol. 59, 1201 – 1206. Maggioni L., Phillip M., Andersen B. Diederichsen A. 2001. Cruciferae: Brassica. and Jørgensen R. 2011. AFLP analysis In: Hanelt P. and IPK (eds): Mansfeld’s of genetic diversity in leafy kale Encyclopedia of Agricultural and (Brassica oleracea L. convar. acephala Horticultural Crops, Vol 3, pp. 1435 - (DC.) Alef.) landraces , cultivars and 1465. Springer, Berlin. wild populations in Europe. Genet. Diederichsen A. and McVetty P.B.E. 2011. Resourc. Crop Evol. 58, 657 – 666. Botany and Plant Breeding, pp. 29 – 56. Clauss E. 1975. Methoden der In: Daun et al. (2011). Artbastardierung innerhalb der Gattung Downey R.K. and Rakow G.F.W. 1987. Brassica zur Schaffung neuen Rapeseed and mustard. In: Fehr W.R. Ausgangsmaterials für die Züchtung. (ed.) Principles of cultivar development, Tag.-Ber. Akad. Landwirtsch. DDR 145, volume 2, crop species. Macmillan, New 83 – 98. York, pp. 437 - 486. Cleemput S. and Becker H.C. 2012. Genetic Downey R.K. and Röbbelen G. 1989. variation in leaf and stem glucosinolates Brassica species. In: Röbbelen G., in resynthesized lines of winter rapeseed Downey R.K. and Ashri A. (eds), Oil (Brassica napus L.). Genet. Resour. Crops of the World, pp. 339 – 362. Crop Evol. 59, 539 – 546. McGraw-Hill, New York. Coulter F.C. 1941. The story of the garden Farnham M.W., Davis E.H., Morgan J.T. vegetables. V. Cabbage: Planted the and Smith J.P. 2008. Neglected world over for peasant and king. Seed landraces of collards (Brassica oleracea World, 18 – 19. L. var. viridis) from the Carolinas

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 472

(USA). Genet. Resour. Crop Evol. 55, Gladis T. and Hammer K. 1990. Die 797 – 801. Gaterslebener Brassica Kollektion – eine Fiori A. 1923/25. Nuova Flora Analitica Einführung. Kulturpflanze 38, 121 - 156. d’Italia. Firenze. Gladis T. and Hammer K. 1992. Die Frandsen H.N. and Winge Ö. 1932. Brassica Gaterslebener Brassica-Kollektion – napocampestris, a new constant Brassica juncea, B. napus, B. nigra und amphidiploid species hybrid. Hereditas B. rapa. Feddes Rep. 103, 469 – 507. 16, 212 – 218. Gladis T. and Hammer K. 1994. Fu J., Zhang M.-F. and Qi X.-H. 2006. Evolutionary classification of the Genetic diversity of traditional Chinese Brassica oleracea group. ISHS Mustard crops Brassica juncea as Symposium on Brassicas, Ninth Crucifer revealed by phenotypic differences and Genetics Workshop. Abstracts of oral RAPD markers. Genet. Resour. Crop papers and posters, p. 55. Evol. 53, 15 – 1519. Gladis T. and Hammer K. 2001. Gates R.R. 1953. Wild cabbages and the Nomenclatural notes on the Brassica effects of cultivation. J. Genetics 51, 363 oleracea-group. Genet. Resour. Crop – 372. Evol. 48, 7-11. Gepts P. 2002. A comparison between crop Gladis T. and Hammer K. 2002. The domestication, classical plant breeding relevance of plant genetic resources in and genetic engineering. Crop Sci. 42, plant breeding. FAL Agriculture 1780 – 1790. Research, Special issue 228, 3 – 13. Gepts P. 2004. Crop domestication as long- Gladis T. and Hammer K. 2003. Die term selection experiment. Plant Brassica-oleracea-Gruppe. Schriften des Breeding Reviews 24, 1 – 44. Vereins zur Erhaltung der Gepts P. and Papa R. 2003. Possible effects Nutzpflanzenvielfalt 1, 79 pp. + of (trans)gene flow from crops on the Appendix. VEN, Lennestadt. genetic diversity of landraces and their Gómez-Campo C. 2005. Assessing the wild relatives. Environ. Biosafety Res. contribution of genebanks: the case of 2, 89 – 103. the UPM seed bank in Madrid. Plant Girke A., Schierold A. and Becker H.C. Genetic Resources Newsletter 151, 33 - 2012. Extending the rapeseed genepool 42. with resynthesized Brassica napus L. I: Gómez-Campo C. and Prakash S. 1999. Genetic diversity. Genet. Resour. Crop Origin and Domestications. In: Gómez- Evol. 59, 1441 – 1447. Campo, C. (ed.) Developments in Plant Gladis T. 1989. Die Nutzung einheimischer Genetics and Breeding, 4. Biology of Insekten (Hymenopteren und Dipteren) Brassica Coenospecies. Elsevier, zur Bestäubung von Kulturpflanzen in Amsterdam, pp. 33 - 58. der Genbank Gatersleben. Kulturpflanze Gómez-Campo C., Aguinagalde I., 37, 79 – 126. Ceresuela J.L., Lázaro A., Martínez- Gladis T. 1995. Crossing experiments in Laborde J.B., Parra-Quijano M., cultivated Brassica oleracea. PGR Simonetti E., Torres E. and Tortosa M.E. Newsletter 104, 32. 2005. An exploration of wild Brassica Gladis T. and Hammer K. 1994. The oleracea L. germplasm in northern Gatersleben Brassica collection, an Spain. Genet. Resour. Crop Evol. 52, 7 - actualized survey. Curcif. Newsl. 16, 6. 13.

473 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Gustafsson M. and Lannér-Herrera C. 1997. EUCARPIA Genetic Resources Section Overview of the Brassica oleracea Meeting, Piešt’any, Slovak Republic, 23 complex: their distribution and May – 26 May 2007. ecological specifications. In: Valdés B. Hammer K., Knüpffer H., Laghetti G. and et al. (eds), Proc. Workshop Cons. Wild Perrino P. 1992. Seeds from the Past. A Relatives , pp. 27 – 32. Palermo. Catalogue of Crop Germplasm in South Hammer K. 1984. Das Italy and Sicily. C.N.R., Bari. Domestikationssyndrom. Kulturpflanze Hammer K., Knüpffer H., Laghetti G. and 32, 11 – 34. Perrino P. 1999. Seeds from the Past. A Hammer K. 1998. Genepools – structure, Catalogue of Crop Germplasm in availability and elaboration for breeding Central and North Italy. C.N.R., Bari. (Germ., Engl. Summary). Schriften Gen. Hammer K., Knüpffer H., Xhuveli L. and Res. 8, 4 – 14. Perrino P. 1996. Estimating genetic Hammer K. 2003. Kulturpflanzenevolution erosion in landraces – two case studies. und Biodiversität. Nova Acta Genet. Resour. Crop Evol. 43, 329 – Leopoldina NF 87, 133 – 146. 336. Hammer K. and Diederichsen A. 2009. Hammer K., Laghetti G. and Perrino P. Evolution, status and perspectives for 1989. Collection of plant genetic landraces in Europe, pp. 23 – 44. In: M. resources in South Italy, 1988. Veteläinen, V. Negri and N. Maxted Kulturpflanze 37, 401 – 414. (eds). European landraces : on farm Hammer K., Laghetti G. and Pignone D. conservation, management and use. (eds) 2011. Linguistic Islands and Plant Bioversity Technical Bulletin No. 15, Genetic Resources – The case of the 344 pp. Arbëreshë. Aracne, Roma, 337 pp. Hammer K. and Laghetti G. 2005. Genetic Hammer K., Laghetti G., Cifarelli S. and erosion – examples from Italy. Genet. Perrino P. 1990. Collection of plant Resour. Crop Evol. 52, 629 – 634. genetic resources in Italy 1989. Hammer K. and Perrino P. 1985. A check- Kulturpflanze 38, 311 – 323. list of the cultivated plants of the Ghat Hanelt P. 1986a. Cruciferae, pp. 272 – 332. oases. Kulturpflanze 33, 269 – 286. In: Schultze-Motel J. (Hrsg.), Rudolf Hammer K. and Perrino P. 1985. A check- Mansfelds Verzeichnis list of the cultivated plants of the Ghat landwirtschaftlicher und gärtnerischer oases. Kulturpflanze 33, 269 – 286. Kulturpflanzen (ohne Zierpflanzen), 2. Hammer K., Esquivel M. and Knüpffer H. Aufl. Akademie-Verlag, Berlin. (eds) 1992 – 1994. “…y tienen faxones y Hanelt P. 1986b. Formal and informal fabas muy diversos de los nuestros …” classifications of the infraspecific Origin, Evolution and Diversity of variability of cultivated plants – Cuban Plant Genetic Resources. 3 vols., advantages and limitations. In: Styles, 824 pp. IPK, Gatersleben. B.T. (ed.), Infraspecific Classification of Hammer K., Kang J.-H. and Laghetti G. Wild and Cultivated Plants, p. 139 – 2007. Wild gathered plants and 156. Oxford. plant domestication – case studies of two Hanelt P. and Hammer K. 1987. Einige distant areas (Italy and Korea). Plant infraspezifische Umkombinationen und Genetic Resources and their Exploitation Neubeschreibungen bei Kultursippen in Plant Breeding for Food and von Brassica L. und Papaver L. Feddes Agriculture. Book of Abstracts, 18th Rep. 98, 553 – 555.

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 474

Hansen L.B., Siegismund H.R. and Helm J. 1963c. Die „Chinakohle“ im Jørgensen R.B. 2001. Introgression Sortiment Gatersleben iii., 3. Brassica between oilseed rape (Brassica napus narinosa L.H. Bailey. Kulturpflanze 11, L.) and its weedy relative, B. rapa L. in 416 – 421. a natural population. Genet. Resour. Helm J. 1964. Unpubl. Die “Chinakohle” im Crop Evol. 48, 621 – 627. Sortiment Gatersleben IV, 4. Brassica Harberd D.J. 1972 A contribution to the alboglabra L.H. Bailey. Published under cyto-taxonomy of Brassica (Cruciferae) Gladis and Hammer (2003). and its allies. Bot. J. Linn. Soc. 65, 1 - Hetterscheid W.L.A. and Brandenburg W.A. 23. 1995. Culton versus taxon: conceptual Harlan J.R. and de Wet J.M J. 1971. Toward issues in cultivated plant systematics. a rational classification of cultivated Taxon 44, 161 – 175. plants. Taxon 20, 509 - 517. Hew D.V.P. and Rumball N. 2000. Walking Hedge I.C. 1976. A systematic and sticks as seed savers – the case of the geographical survey of the old world Jersey kale (Brassica oleracea L. Cruciferae. In: Vaughan J.C., MacLeod convar. acephala (DC.) Alef. var. viridis A.J. and Jones B.M.G. (eds) The L.). Econ. Bot. 54, 141 – 143. Biology and Chemistry of the Hoang H.-Dz., Knüpffer H. and Hammer K. Cruciferae. Academic Press, New York, 1997. Additional notes to the checklist pp. 1 - 45. or Korean cultivated plants (5). Helm J. 1959. Brassica oleracea L. In: Consolidated summary and indexes. Mansfeld R., Vorläufiges Verzeichnis Genet. Resour. Crop Evol. 44, 349 – landwirtschaftlich oder gärtnerisch 391. kultivierter Pflanzenarten. Kulturpflanze, Hodgkin T. 1995. Cabbages, kale, etc.In: Beihfeft 2, 78 – 84. Smartt J. and Simmonds N.W. (eds), Helm J. 1960. Brokkoli und Spargelkohl. Evolution of Crop Plants, 2nd ed., pp. 76 Beiträge zur Geschichte ihrer Kultur und – 82. Longman, London. zur Klärung ihrer morphologischen und Hondelmann W. 2002. Die Kulturpflanzen taxonomischen Beziehungen der griechisch-römischen Welt. untereinander sowie zum Blumenkohl. Pflanzliche Ressourcen der Antike. Züchter 30, 223 – 241. Gebrüder Bornträger Berlin. Helm J. 1961. Die “Chinakohle” im Hu Q., Li Y. and Mei D. 2009. Introgression Sortiment Gatersleben I., 1. Brassica of genes from wild crucifers. In: Gupta pekinensis (Lour.) Rupr. Kulturpflanze S.K. (ed), Biology and Breeding of 9, 88 – 113. Crucifers, pp. 261 – 283. CRC Press, Helm J. 1962. Die laciniaten Sippen von Boca Raton. Brassica oleracea L. Kultrurpflanze 10, Hu Z.-Y., Hua W., Huang S.-M. and Wang 111 – 121. H.-Z. 2011. Complete chloroplast Helm J. 1963a. Morphologisch- genome sequence of rapeseed (Brassica taxonomische Gliederung der napus L.) and its evolutionary Kultursippen von Brassica oleracea L. implications. Genet. Resour. Crop Evol. Kulturpflanze 11, 92 - 210. 58, 875 – 887. Helm J. 1963b. Die „Chinakohle“ im Huangfu C.-H., Song X.-L. and Quiang S. Sortiment Gatersleben II., 2. Brassica 2009. ISSR variation within and among chinensis Juslen. Kulturpflanze 11, 333 wild Brassica juncea populations: – 357. implication for herbicide resistance

475 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

evolution. Genet. Resour. Crop Evol. 56, Larkcom J. 1987. Chinese Brassicas – in 913 – 924. China. The Garden 112, 325 – 330. Johnston J.S., Pepper A.E., Hall A.E., Chen Lee S.-H. 1982. Vegetable crops growing in Z.J., Hodnett G., Drabek J., Lopez R. China. Sci. Hort. 17, 201 – 209. and Price H.J. 2005. Evolution and Li C.-Y., Zhang G.-Y., Hammer K., Yang genome size in Brassicaceae. Annals of C.-Y. and Long C.-L. 2011. A checklist Botany 95, 229 - 235. of the cultivated plants Yunnan (PR Khoshbakht K. and Hammer K. 2010. China). Genet. Resour. Crop Evol. 58, Threatened Crop Species Diversity. 153 – 164 + electr. appendix. Shahid Beheshti Univ. Press, Lin W. 1980. A study on the classification Teheran, 134 pp. of Chinese cabbages. Acta Hort. Sincia 7 Kim H.-S., Hammer K., Han U.-X., Hanelt (2), 21 – 28. P. and Pak H.-S. 1987. Missions for the Lin Y and Lee S.H. 1985. A new botanical collection of plant genetic resources in variety of stem mustard Baoercai. Acta the Democratic People’s Republic of Hort. Sinica 12, 41 – 44. Korea 1986 for the collection of Linnaeus C. 1753. II, ed. landraces of cultivated plants. 1. Holmiae. Kulturpflanze 35, 355 – 365. Linnaeus C. 1763. Species plantarum, ed. 2. Kimber D.S. and McGregor D.I. 1995. The Holmiae. species and their origin, cultivation and Liu A.-H. and Wang J.-B. 2006. Genomic world production. In: Kimber D. and and evolution of Brassica allopolyploids McGregor D.I. (eds), Brassica Oilseeds: revealed by ISSR markers. Genet. Production and Utilization. CABI, Resour. Crop Evol. 53, 603 – 611. Wallingford, pp. 1 – 9. Louarn S., Torp A.M., Holme I.B., Kitamura S. 1950. The cultivated Brassicae Andersen S.B. and Jensen B.D. 2007. of China and Japan. Mem. Coll. Sci. Database derived microsatellite markers Univ. Kyoto, ser. B 19, 75 – 80. (SRs) for cultivar differentiation in Laghetti G., Martignano F., Falco V., Brassica oleracea. Genet. Resour. Crop Cifarelli S., Gladis T. and Hammer K. Evol. 54, 1717 – 1725. 2005. “Mugnoli”: a neglected race of Maffi L. (ed.) 2001. On Biocultural Brassica oleracea L. from Salento Diversity. Linking language, culture and (Italy). Genet. Ressour. Crop Evol. 52, the environment. Smiths. Inst. Press, 635 – 639. Washington. Laghetti G., Pignone D., Cifarelli S., Maggioni L., von Bothmer R., G. Poulsen Martignano F., Falco V., Traclò B.R.G. and Branca F. 2010. Origin and and Hammer K. 2008. Agricultural domestication of cole crops (Brassica in Grecia and Bovesia, the oleracea L.): linguistic and literary two Griko-speaking areas in Italy. Plant considerations. Econ. Bot. 64, 109 – Gen. Res. Newsletter 156, 43 – 49. 123. Landsrath S. und Hammer K. 2007. Makino T. 1912. Yokusai Iinuma: Somoku- Pflanzliche Agrarbiodiversität – eine Dzusetsu; or an iconography of plants essayistische Überarbeitung Friedrich indigenous to, cultivated in, or Alefelds „Landwirthschaftlicher Flora“ introduced into Nippon (Japan.). 3rd ed., von 1866. Schriften des VEN 6, P.1, Herbaceous Plants. Cremlingen – Schandelah, 164 pp. Maly R., Hammer K. and Lehmann Chr. O. 1987. Sammlung pflanzlicher

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 476

genetischer Ressourcen in Süditalien – Metzger J. 1833. Systematische ein Reisebericht aus dem Jahre 1950 mit Beschreibung der kultivierten Kohlarten. Bemerkungen zum Schicksal der Heidelberg. Landsorten “in situ” und in der Mithen R.F., B.G. Lewis, R.K.Cheaney and Genbank. Kulturpflanze 35, 109 – 134. G.R. Fenwick 1987. Glucosinolates of Mansfeld R. 1953. Zur allgemeinen wild and cultivated Brassica species. Systematik von Kulturpflanzen 1. Phytochemistry 26, 1969 – 1973. Kulturpflanze 1, 138 – 155. Mizushima U. 1980. Genome analysis in Mansfeld R. 1954. Zur allgemeinen Brassica and allied genera. In: Tsunoda Systematik von Kulturpflanzen 2. S., Hinata K. and Gómez-Campo C. Kulturpflanze 2, 130 – 142. Brassica crops and wild allies – biology Mansfeld R. 1959. Vorläufiges Verzeichnis and breeding. Japan Scientific Societies landwirtschaftlich und gärtnerisch Press, Tokyo, pp. 89 - 106. kultivierter Pflanzenarten (mit Murphy D.J. 2007. People, Plants & Genes. Ausschluß von Zierpflanzen). Oxford University Press, New York ,401 Kulturpflanze Beih. 2, 659 pp. pp. Maxted N., Ford-Lloyd B.V., Kell S.P., Olsson G. 1954. Crosses within the Iriondo J., Dulloo E. and Turok J. 2008. campestris group of the genus Brassica. Conservation and Heriditas 40, 398 – 418. Use. CAB International, Wallingford. Onno M. 1933. Die Wildformen aus dem McGrath J.M. and Quiros C.F. 1992. Verwandtschaftskreis “Brassica Genetic diversity at isozyme and RFLP oleracea L.” Österr. Bot. Z. 82, 309 – loci in Brassica campestris as related to 334. crop type and geographical origin. TAG Osborn T.C., Pires J.C., Birchler J.A., Auger 83, 783 – 790. D.L., Chen Z.J., Lee H.S. et al. 2003. McNeill J., Barrie F.R., Burdet H.M., Understanding mechanisms of novel Demoulin V., Hawksworth D.L., gene expression in polyploids. Trends in Marhold K., Nicolson D.H., Prado J., Genetics 19, 141 – 147. Silva P.C., Skog J.E., Wiersema J.H. and Palmer J.D., Shields C.R., Cohen D.B. and Turland N.J. (eds) 2006. International Orton T.J. 1983. Chloroplast DNA Code of Botanical Nomenclature evolution and the origin of amphidiploid (Vienna Code). Adopted by the Brassica species. Theoretical and Seventeenth International Botanical Applied Genetics 65, 181-189. Congress, Vienna, Austria, July 2005. Perrino P. and Hammer K. 1985. Collection Regnum Vegetabile 146. A.R.G. of land-races of cultivated plants in Gantner Verlag KG, Ruggell, South Italy, 1984. Kulturpflanze 33, 225 Liechtenstein. – 236. Mei J., Li Q., Yang X, Quian L., Lin L., Yin Perrino P., Pignone D. and Hammer K. J., Frauen M., Li J. and Quian W. 2010. 1992. The occurrence of wild Brassica Genomic relationships between wild and of the the oleracea group (2n – 18) in cultivated Brassica oleracea L. with Calabria (Italy). Euphytica 59, 99 – 101. emphasis on the origination of cultivated Pickersgill B. 2009. Domestication of plants crops. Genet. Resour. Crop Evol. 57, revisited – Darwin to the present day. 687 – 692. Bot. J. Linn. Soc. 161, 203 – 212. Pickersgill B. and Karamura D.A. 1999. Issues and options in the classification of

477 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

cultivated bananas, with particular Rakow G. 2004. Species origin and reference to East African highland economic importance of Brassica. In: bananas. In: Andrews S. et al. (eds), pp. Pua E.C. and Douglas C.J. (eds.) 159 – 164. Biotechnology in Agriculture and Pignone D. and Gómez Campo C. 2010. Forestry. Springer, Berlin – Heidelberg, Eruca. In: Cole Ch. et al. (ed), Wealth of pp. 3-11. Wild Species: Role in Plant Genome Reiner H., Holzner W. and Ebermann R. Elucidation and Improvement. Elsevier. 1995. The development of turnip-type Pistorius R. 1997. Scientists, Plants and and oilseed types Brassica rapa crops Politics. A History of Plant Genetic from wild-type in Europe – an overview Resources Movement. IPGRI, Rome. of botanical, historical and linguistic Prakash S., Takahata Y., Kirti P.B. and facts. Rapeseed Today Tomorrow 4, Chopra V.L. 1999. Cytogenetics. In: 1066 – 1069. Gómez-Campo, C. (ed) Developments in Reiner L., Gladis Th., Amon H. and plant genetics and breeding, 4. Biology Emmerling-Skala A. 2005: The of Brassica coenospecies. Elsevier, ‘Bavarian Turnip’ – a rediscovered local Amsterdam, pp. 59 - 106. vegetable variety of Brassica rapa L. Proskowetz E. von and Schindler F. 1890. em. Metzg. var. rapa. Genetic Res. and Welches Werthverhältnis bestgeht Crop Evol. 52, 111-113. zwischen den Landrassen Röbbelen G. 1960. Beiträge zur Analyse des landwirthschaftlicher Kulturpflanzen Brassica Genoms. Chromosoma 11, 205 und den sogenannten Züchtungsrassen? - 228. Internationaler Land- und Sabnis T. and Phatak M.G. 1935. A Forstwirthschaftlicher Kongress zu Wien preliminary note on the classification of 1890, Heft 13, pp. 3 – 24. cultivated Indian mustards. Indian Agric. Purugganan M.D. and Fuller D.Q. 2009. The Sci. 5, 559 – 578. nature of selection during plant Sánchez-Yélamo M.D. 2009. Relationships domestication. Nature 457, 843 – 848. in the Diplotaxis-Erucastrum-Brassica Qi X.-H., Yang J.-H. and Zhang M.-F. 2008. complex (Brassicaceae) evaluated from AFLP-based genetic diversity isoenymatic profiles of the accessions as assessment among Chinese vegetable a whole. Applications for mustards (Brassica juncea (L.) Czern.). characterisation of phylogenetic Genet. Resour. Crop Evol. 55, 705 – resources preserved ex situ. Genet. 711. Resour. Crop Evol. 56, 1023 – 1036. Qi X.-H., Zhang M.-F. and Yang J.-H. 2007. Šebalina M.A. 1968. Istorija botaničeskogo Molecular phylogeny of Chinese izučenii i klassfikacii repy. Trudy prikl. vegetable mustard (Brassica juncea) bot., gen. i sel. 38, 44 – 87. based on the internal transcribed spacers Šebalina M.A. and Sazonova L.V. 1985. (ITS) of nuclear ribosomal DNA. Genet. Root crops (Brassica – turnip, rutabaga, Resour. Crop Evol. 54, 1709 – 1716. ). In: Dorofeev V.F. (ed.), Flora of Rabbani M.A., Aki I., Yoshie M., Tohru S. the Cultivated Plants (USSR) vol. 18. and Kenji T. 1998. Genetic diversity in Leningrad. mustard (Brassica juncea L.) germplasm Seyis F., Snowdon R.J., Lühs W. and Friedt from Pakistan as determined by RAPDs. W. 2003. Molecular characterization of Euphytica 103, 235 – 242. novel resynthesized rapeseed (Brassica napus) lines and analysis of their genetic

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 478

diversity in comparison with spring Encyclopedia of Agricultural and rapeseed cultivars. Plant Breeding 122, Horticultural Crops 3. Springer, Berlin. 473 - 478. Sun V.G. 1946. The evolution of taxonomic Sheng X., Wen G., Guo Y., Yan, H., Zhao characters of cultivated Brassica with a H. and Liu F. 2012. A semi-fertile key to species and varieties. 1. The interspecific hybrid of Brassica rapa and characters; The key. Bull. Torrey Bot. B. nigra and the cytogentic analysis of Club 73, 244 – 281, 370 – 377. its progeny. Genet. Resour. Crop Evol. Takuno S., Kawahara T. and Ohnishi O. 59, 73 – 81. 2007. Phylogenetic relationships among Sinskaja E.N. 1928. The oleiferous plants cultivated types of Brassica rapa L. em. and root crops of the familiy Cruciferae Metzg. as revealed by AFLP analysis. (Russ., Engl. summary). Bulletin for Genet. Resour. Crop Evol. 54, 279 – Applied Botany, Genetics and Plant 285. Breeding 19, 3 - 648. Toxopeus H. and Oost E.H. 1985. A cultivar Sinskaja E.N. 1969. Istoričeskaja geografia group classification of Brassica rapa L. kul’turnoj flory. Kolos, Leningrad. Cruciferae Newsl. 10, 6 – 7. Snogerup S. 1980. The wild forms of the Tsen M. and Lee S.-H. 1942. A preliminary Brassica oleracea group (2n=18) and study of cultivated Brassica. Hortus their possible relations to the cultivated Sinicus, Bull. 2, Chunking pp. 1 – 31. ones. In: Tsonuda S., Hinata K. and Tsunoda S., Hinata K. and Gómez-Campo Gómez-Campo C. (eds.) Brassica crops C. (eds) 1980. Brassica crops and wild and wild allies biology and breeding. allies: biology and breeding. Sci. Soc. Japan. Sci Soc. Press, pp. 121 - 132. Press, Tokyo. Snogerup S., Gustafsson M. and von U N. 1935. Genomic analysis in Brassica Bothmer R. 1980. Brassica sect. with special reference to the Brassica (Brassicaceae) I. Taxonomy experimental formation of B. napus and and variation. Willdenowia 19, 271 - peculiar mode of fertilization. Japanese 365. Journal of Botany 7, 389 - 452. Snowdon R.J. 2007. Cytogenetics and Van Treuren R. and Bas N. 2008. Perennial genome analysis in Brassica crops. kales: collection rationalization and Chromosome Research 15, 85 - 95. genetic relatedness to other Brassica Song K. and Osborn T.C. 1992. Polyphyletic oleracea crop types. Genet. Resour.Crop origins of Brassica napus: new evidence Evol.. 55, 203 – 210. based on organelle and molecular RFLP Vaughan D.A., Balász E. and Heslop- analyses. Genome 35, 992 - 1001. Harrison J.S. 2007. From crop Song K., Osborne T.C. and Williams P.H. domestication to super-domestication. 1990. Brassica taxonomy based on Ann. Bot. 100, 893 – 901. nuclear restriction fragment length Vellvé R. 1993. The decline of diversity in polymorphism (RFLPs): 3. Genome European agriculture. Ecologist 23, 64 – relationships in Brassica and related 69. genera and the origin of B. oleracea and Warwick S.I., Francis A. and Gugel R.K. B. rapa (syn. B. campestris). Theor. 2009. Guide to Wild Germplasm, Appl. Genet. 79, 497 – 506. Brassica and allied crops (tribe Specht C.E. 2001. Cruciferae Brassiceae, Brassicaceae). 3rd (Brassicaceae), pp. 1413 – 1481. In: edition.http://www.brassica.info/info/pu Hanelt P. and IPK (eds), Mansfeld’s blications/guide-wild-germplasm.php

479 International Journal of AgriScience Vol. 3(6): 453-480, June 2013

Warwick S.I., Gugel R.K., McDonald T. and determined by RAPD analysis? Genet. Falk K.C. 2006: Genetic variation in Resour. Crop Evol. 45, 105 – 111. Ethiopian mustard (Brassica carinata A. Zeven A.C. 1998. Landraces: a review of Braun) germplasm in western Canada. definitions and classifications. Euphytica Genet. Resour. Crop Evol. 53, 297 – 104, 127 – 139. 312. Zhou W.J., Zhang G.C., Tuvesson S., Warwick S.I., James T. and Falk K. 2008. Dayteg C. and Gertsson B. 2006: AFLP-based molecular characterization Genetic survey of Chinese and Swedish of Brassica rapa and diversity in oilseed rape (Brassica napus L.) by Canadian spring turnip rape cultivars. simple sequence repeats (SSRs). Genet. Plant Genetic Resources Resour. Crop Evol. 53, 443 – 447. Characterization and Utilization 6, 11 - Zhukovsky P.M. 1933. La Turquie agricole. 21. Moskva-Leningrad. Wu X.-M., Chen B.-Y., Lu G., Wang H.-Z., Zohary D., Hopf M. and Weiss E. 2012. Xu K., Guizhan G. and Song Y. 2009. Domestication of Plants in the Old Genetic diversity in oil and vegetable World. 4th ed., Oxford University Press, mustard (Brassica juncea) landraces Oxford. revealed by SRAP markers. Genet. Resour. Crop Evol. 56, 1011 – 1022. Yamagishi H. and Takayanagi K., 1982. Cross-compatibility of hakuran (artificially synthesized Brassica napus) with Brassica vegetables. Cruciferae Newsletter 7: 34-35. Yang B. and Quiros C.F. 2010. Survey of glucosinolate variation in leaves of Brassica rapa crops. Genet. Resour. Crop Evol. 57, 1079 – 1089. Yang Y.W., Lai K.N., Tai P.Y., Ma D.P. and Li W.H. 1999 Molecular phylogenetic studies of Brassica, , and allied genera based on the internal transcribed spacer region of 18S-25S rDNA. and Evolution 13, 455 - 462. Yang-Zhen L. 1986. Chromosome numbers in Brassiceae (Cruciferae) in China. Acta Phytotax. Sin. 24, 268 – 272. Yarnell S.H. 1956. Cytogenetics of the vegetable crops. II. Crucifers. Botanical Review 22, 81 - 166. Zeven A., Dehmer K., Gladis Th., Hammer K. and Lux H. 1998. Are duplicates of perennial kale (Brassica oleracea L. var. ramosa DC.) true duplicates as

International Journal of AgriScience Vol. 3(6): 453-480, June 2013 480