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• W2102952994 abstract "Domestication is generally seen as a process of human-mediated selection and diffusion of an organism to provide food and/or other valuable commodities. It is an open-ended process, with diffuse beginnings and no known endpoint, during which the artificial selection of domesticates is enhanced by gene exchange with their wild relatives (Darwin, 1868). Genetic knowledge of the process of domestication is important from a cultural and an agricultural standpoint. It tells us about the origin and history of human civilizations and can inform the management of cattle and crop genetic resources (Doebley et al., 2006; Larson et al., 2014). The effects of introgression by secondary contacts is particularly relevant at a local scale because gene flow between domesticates and their wild relatives can facilitate secondary diversification of crops and livestock, and can certainly contribute to the emergence of new local varieties or races. It has recently been shown that domestication of the grape or the apple is in part a history of repeated introgressions with wild relatives, and that the diffusion of these crops has even led to the selection in Western Europe of varieties that are closer genetically to their secondary progenitors (Myles et al., 2011; Cornille et al., 2012). However, making the distinction between single and multiple independent primary domestications is often hard and the data necessary to substantiate such a claim is not always available. In a recent study published in New Phytologist, Díez et al. (2015) argue for at least two independent domestication events of the olive tree, opposing the predominant view of a single primary domestication followed by several secondary diversification events. The olive (Olea europaea subsp. europaea; Box 1) is often considered the most iconic tree of the Mediterranean world. The wild olive (var. sylvestris) occurs around the Mediterranean Basin and the first signs of its domestication accompanied the emergence of some of the most ancient civilizations (Loumou & Giourga, 2003; Kaniewski et al., 2012). In historical times, the cultivated olive (var. europaea) has been primarily a source of oil, although the original use of this oil remains a matter of debate. The most primitive descriptions of the use of olive oil seem to indicate that it was a combustible to produce light and a body ointment of ritual significance. For instance, olive oil was burned in Moses' menorah according to the book of Exodus in the Ancient Testament. Conversely, the culinary use of the oil is not widely reported until the ancient Roman epoch (Tardi, 2014). The olive tree is also a source of wood, and its leaves are palatable to cattle and have been widely used as forage (already mentioned during the third century ad by Claudius Aelianus in Natura Animalium). Six wild olive subspecies are recognized (Table 1) and usually considered as primary genetic resources for the cultivated olive, although natural hybridization has been reported only between diploids (Zohary, 1994; Green, 2002; Besnard et al., 2013b, 2014). The main wild progenitor of the cultivated olive (O. e. subsp. europaea var. europaea) is clearly the wild Mediterranean olive or oleaster (O. e. subsp. europaea var. sylvestris). The two have overlapping distributions in the Mediterranean Basin, often occurring in sympatry, share similar ecological requirements and are morphologically very similar. The distinction is usually done based on fruit size, more specifically on the thickness of the mesocarp. Other characteristics that have been used in the past to discriminate between both forms such as the leaf size and shape, the architecture, or the presence of spines are plastic and under environmental rather than genetic control (Green, 2002). Many studies have tried to establish a clear genetic distinction between varieties europaea and sylvestris. As a result, high differentiation between cultivated olive and supposedly genuinely wild olives has been reported in the western Mediterranean Basin (Besnard et al., 2001; Lumaret et al., 2004), especially in Corsica (Bronzini de Caraffa et al., 2002), Morocco (Lumaret et al., 2004), Tunisia (Hannachi et al., 2010), Italy (Baldoni et al., 2006), and Spain (Belaj et al., 2010). These same genetic studies also indicate frequent admixture between different genetic pools, partly as a consequence of the cultivated olive diffusion from the easternmost to westernmost parts of the Mediterranean Basin and its subsequent cross with local oleaster populations. Over the last few years many authors have been accumulating palaeobotanical, archaeological, historical, and molecular data on the biogeography of the wild olive and the history of its domestication (Terral et al., 2004; Baldoni et al., 2006; Carrión et al., 2010; Kaniewski et al., 2012; Besnard et al., 2013a). Multiple origins of cultivars have been demonstrated and three genetic pools independently identified by different research teams (Haouane et al., 2011; Belaj et al., 2012; Díez et al., 2012). Given that the olive tree may have persisted around the Mediterranean Basin as part of the natural vegetation since the Late Tertiary (Palamarev, 1989), the question remains as to whether these groups of cultivated olives derived from a single initial domestication event in the Levant followed by secondary diversification or whether they are the result of independent domestication events. Some recent analyses suggested that the domestication of the olive tree primarily occurred in the Levant followed by multiple, secondary diversifications across the Mediterranean (Besnard et al., 2013a,b). In their new paper, Díez et al. (2015) compare scenarios of diffusion and admixture between different genetic pools using an approximate Bayesian computation (ABC) framework (Beaumont, 2010). This approach is powerful and can provide valuable insights to the analysis of domestication patterns. ABC modeling has been recently applied to decipher the domestication history of other fruit trees (see Gerbault et al., 2014, for a review) but it had not been used before to study the cultivated olive origins. Using this modeling framework to analyze nuclear genomic data, Díez et al. (2015) show three genetic pools in the eastern, central and western Mediterranean. According to their results, these three pools resulted from a complex admixture between eastern and western cultivars and oleasters. Their results also support a relatively recent origin of south Iberian cultivated varieties (group Q1), likely following a genetic bottleneck. With co-ancestry analyses, they identified two ancient varieties that could be the main Q1 ancestors, similar to what has been shown for the local selection of grapes (Bowers et al., 1999). Based on their results, Díez et al. (2015) propose two alternative scenarios of domestication. The first hypothesis that they consider is a single primary domestication in the eastern Mediterranean, followed by human-mediated spread of the crop and secondary diversifications, in agreement with the predominant view in the field and the results of other recent studies (Besnard et al., 2013a,b). The second hypothesis, which Díez et al. (2015) favor, is that of two independent primary domestication events in eastern and central Mediterranean. In light of the data and results presented in the paper we believe there is actually no statistical support to favor any of the models. Moreover, the three verbal arguments given by the authors as corroboration for the second hypothesis do not exclude a single primary domestication followed by secondary diversification (Cornille et al., 2012). It is possible that the reason for these inconclusive results lies in the limited number of wild O. europaea populations used by Díez et al. (2015). To account for the high genetic diversity of the olive tree, a comprehensive sampling incorporating genuinely wild populations of oleasters from across the Mediterranean would be advisable. Díez et al. (2015) analyzed only a few wild populations from Spain, Sardinia and Israel (that include feral or admixed trees). This limited sampling curtails the possibility of testing alternative, complex domestication scenarios. Of course, the fact that the data presented by Díez et al. (2015) are inconclusive does not mean that the olive tree was not primary domesticated in multiple independent events as has been proposed before (Terral, 1997; Besnard et al., 2001). However, this scenario does not fit some of the genetic and historical evidence. Previous phylogeographic studies have shown that two main wild olive genetic pools can be recognized with nuclear genetic markers. These two pools approximately correspond to the western-central and eastern Mediterranean populations (Besnard et al., 2001; Rubio de Casas et al., 2006; Breton et al., 2008). In a recent study that combined a large sample of cultivated olives (470 genotypes) and oleasters (390 genotypes), it was shown that most Mediterranean cultivars were mainly assigned (based on nuclear markers) to the eastern oleaster genetic pool, while no cultivar could be unambiguously assigned to the western one (Besnard et al., 2013b), even those with maternal (i.e. plastid) lineages that originated from the western Mediterranean basin. This result clearly supports that most cultivars either belong to the eastern genetic pool or are admixed forms. Palaeobotanical and archaeological data attest to the human use of O. europaea in the western Mediterranean since the Holocene and the Neolithic period (Terral et al., 2004). Moreover, O. europaea subsp. europaea spread out of Late Glacial Maximum refugia was concomitant with human migration and either humans or the fauna associated with anthropogenic modification of the landscape might have contributed to the dispersal of the olive over the western Mediterranean area, and possibly even into the Saharan mountains (Besnard et al., 2013b). However, it is important to keep in mind that oleasters might have also been used for wood or forage and that there is no unambiguous evidence that fruit traits were successfully selected for and propagated at that point in time (Kaniewski et al., 2012; although see Terral et al., 2004). In other words: to date, there is not enough genetic or archaeological support for an independent primary domestication of the modern olive in the central or western parts of the Mediterranean Basin. Based on the different sources of evidence available, we propose a scenario for post-glacial spread of the wild olive around the Mediterranean Basin and for the primary domestication and secondary diversification of the olive tree (Fig. 1). As proposed by Díez et al. (2015), this framework could be falsified by means of ABC modeling. In particular, we could test for an initial bottleneck during primary domestication in the eastern Mediterranean. However, any explicit model should build on previous biogeographic and archaeological knowledge and will require appropriate sampling including genuinely wild olives that still persist in northern Levant and Cyprus (Besnard et al., 2013a). To clearly identify the area of primary domestication and secondary diversification, the model should consider alternative parameterizations of the contribution of the eastern and western oleaster genetic pools to the extant stock. In light of the genetic diversity of both the crop and the wild olive, this is only possible if the model fits a dataset that includes a wide sample of wild populations. Additionally, the model should also allow for alternative demographic scenarios so hypotheses concerning the timing of domestication events, and the potential steps from eastern to central Mediterranean and from central to western Mediterranean could be explicitly parameterized. Given the cultural and economic relevance of this tree, characterizing the origin and history of the olive is a worthy effort that should be undertaken without leaving out any relevant data or explanatory hypotheses. The authors thank Pablo Vargas, Lounes Chikhi, Christophe Thébaud and two anonymous referees for their helpful comments. G.B. is supported by TULIP (ANR-10-LABX-0041)." @default.
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• W2102952994 title "Single vs multiple independent olive domestications: the jury is (still) out" @default.
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