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(Spinacia Oleracea L.) the Possible Origin and Spread of Spinach

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(Spinacia Oleracea L.) the Possible Origin and Spread of Spinach A review on the domestication and breeding history of spinach (Spinacia oleracea L.) The possible origin and spread of spinach Arnau Ribera Tort MSc Thesis – MPB, Specialization: Molecular Plant Breeding and Pathology PBR-80436 – Reg. Nr. 950418-692-080 22nd May 2019 Supervisors: Prof. Dr. Yuling Bai – Laboratory of Plant Breeding Dr. Chris Kik – Centre for Genetic Resources, the Netherlands Dr. Robbert van Treuren – Centre for Genetic Resources, the Netherlands Dr. Ir. Anne-Marie Wolters – Laboratory of Plant Breeding Summary This thesis consists of two parts, namely “A review on the domestication and breeding history of spinach (Spinacia oleracea L.)” and “The possible origin and spread of spinach”. In the first part, an overview of the crop is presented. Characteristics and relevance of spinach are followed by a brief review of its two wild relatives, the state of its germplasm and its domestication. A subsequent section on spinach breeding history is presented, from what it is available in the first references of the 16th century to current breeding and its future perspectives. In the second part, the phylogenetic relationship and population structure of a selection of spinach landraces and wild spinach accessions have been analysed. The goal of the study is to confirm the phylogenetic relationship between the 3 Spinacia species and study the centre of origin of spinach. Moreover, a selection of landraces from the Eastern Mediterranean and Eastern and Southern Asia has also been included in the analysis to examine if it is possible to relate phylogenetic differences to the hypothetical spread of spinach outside its centre of origin. Index Part I: A review on the domestication and breeding history of spinach (Spinacia oleracea L.) ........ 1 I.1. Importance of spinach .............................................................................................. 1 I.2. Description of spinach ............................................................................................... 1 I.3. Nutritional content and functional properties ................................................................ 2 I.4. Spinach wild relatives ............................................................................................... 4 I.5. State of spinach germplasm ....................................................................................... 6 I.6. Spinach domestication .............................................................................................. 6 I.7. Breeding spinach cultivars: past, present and future .................................................... 10 I.7.1. Disease resistance breeding in spinach .................................................................. 13 I.7.2. Current and future trends in spinach breeding ........................................................ 15 Part II: The possible origin and spread of spinach ................................................................... 20 II.1. Introduction ............................................................................................................ 20 II.2. Materials & methods ................................................................................................ 22 II.3. Results ................................................................................................................... 25 II.4. Discussion .............................................................................................................. 29 II.5. Conclusion .............................................................................................................. 31 References ........................................................................................................................ 33 Appendices ..................................................................................................................... 44 Appendix 1. Morphological details of spinach wild relatives .................................................. 44 Appendix 2. Overview of spinach germplasm ..................................................................... 45 Appendix 2.1. Germplasm of cultivated spinach.............................................................. 45 Appendix 2.2. Germplasm of spinach wild relatives ......................................................... 50 Appendix 3. Breeding history of spinach ............................................................................ 52 Appendix 3.1. Spiny-seeded cultivars ........................................................................... 52 Appendix 3.2. Smooth-leaved smooth-seeded cultivars ................................................... 53 Appendix 3.3. Savoy-leaved smooth-seeded cultivars ..................................................... 55 Appendix 4. Spinacia accessions used in the study .............................................................. 56 Appendix 5. Protocol for DNA isolation using a CTAB buffer .................................................. 58 Appendix 6. SNP information ........................................................................................... 59 Appendix 7. KASP results ................................................................................................ 61 Appendix 8. NEXUS file ................................................................................................... 65 Appendix 9. ΔK analysis for estimating the population structure ........................................... 67 Appendix 10. Population structure analysis for K=1 to 10 .................................................... 68 Part I: A review on the domestication and breeding history of spinach (Spinacia oleracea L.) Spinach [Spinacia oleracea L., 2n=2x=12] is one of the most nutritious consumed vegetables (Morelock and Correll 2008). It is a very versatile crop as it is eaten either raw or cooked, and it is a common ingredient in various cuisines. The history of spinach, both from a domestication and a breeding history perspective, is arguably little known and not many references are available on this topic. For this reason, this study intends to be a review of what is known about spinach and what are the current trends on the breeding of this vegetable. I.1. Importance of spinach The global production of spinach reached more than 26.7 million tonnes in 2016 (FAOSTAT 2018). Especially in China, which is the leading country in spinach production (Table 1), the spinach industry has had a steady growth during the last years. The rest of the top 10 spinach producing regions are presented in Table 1. At a global scale, in 2016 the spinach industry had a value of production of $18 billion and solely China accounted for approximately $15.4 billion of this amount (Table 1). Spinach seed production is mainly based in Denmark (Morelock and Correll 2008; Deleuran 2010; Correll et al. 2011), as this country contains more than 70% of the hectares destined to spinach seed production. The second most important region is the Pacific Northwest of the USA, represented by the states of Washington and Oregon, which accounts for ~ 20% of the seed production land (Du Toit 2018). These two regions share a common mild marine climate with long days to favour flowering (Morelock and Correll 2008). Table 1. Spinach production and value of production of the 10 regions with the largest spinach production. Value of production Global production Global production Region (millions of USD, (%, yr 2016) (%, yr 2000) yr 2016) China 91.50 78.75 15,436.0 European Union 2.21 5.59 464.0 USA 1.36 3.45 371.8 Japan 0.92 3.32 1,172.8 Turkey 0.79 2.15 99.3 Iran 0.44 0.92 27.8 Pakistan 0.40 0.77 17.0 Kenya 0.28 0.14 11.6 South Korea 0.27 1.27 139.2 Malaysia 0.20 0.20 23.1 26.78 9.52 18,013.7 Global million tonnes million tonnes million USD Source: FAOSTAT (2018) I.2. Description of spinach Spinach is a leafy vegetable from the Amaranthaceae family (Hassler 2018). This family from the Caryophyllales order includes other important crops such as beet [Beta vulgaris L.], quinoa [Chenopodium quinoa Willd.] and amaranth [Amaranthus spp.]. Spinach is an annual plant with marked vegetative growth and reproductive phases. Typically, at the end of winter or during early spring seeds are sown and seedlings grow to form a rosette of leaves up to 15 to 30 cm tall. (Krarup and Moreira 1998; Van der Vossen 2004). Many cultivars exist with distinct leaf attributes, from round to hastate shape and from flat to crinkly [savoy] texture (Morelock and Correll 2008). Moreover, the petiole of the leaves is also variable, both its colour and length (Serpolay et al. 2011; Ma et al. 2016). The arrival of warmer and longer summer days induces bolting; hence spinach plants start their reproductive phase by growing a peduncle of about one metre tall, simple or branched 1 (Krarup and Moreira 1998), and generally with terminal staminate flowers and/or pistillate flowers at bract axils (Uotila 1997). Nowadays, there are spinach varieties adapted to different climatic conditions and photoperiods. Some varieties possess bolting resistance to longer and warmer days, which make them compatible with summer cultivation. In a similar fashion, winter-hardy varieties can tolerate colder temperatures and are adapted to winter cultivation (Van der Vossen 2004). This makes spinach a very flexible crop that can be cultivated virtually all year round in many regions. Spinach is a wind-pollinated dioecious species, even though monoecious plants with flowers of both sexes also exist (Khattak et al. 2006). Different
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