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Genetic and Morphological Characterization of the Endangered Austral Papaya Vasconcellea Chilensis (Planch

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Genetic and Morphological Characterization of the Endangered Austral Papaya Vasconcellea Chilensis (Planch Genet Resour Crop Evol (2014) 61:1423–1432 DOI 10.1007/s10722-014-0143-0 NOTES ON NEGLECTED AND UNDERUTILIZED CROPS Genetic and morphological characterization of the endangered Austral papaya Vasconcellea chilensis (Planch. ex A. DC.) Solms B. Carrasco • R. Garcı´a-Gonza´les • C. Dı´az • P. A´ vila • P. Ca´ceres • G. A. Lobos • H. Silva • P. D. S. Caligari Received: 20 January 2014 / Accepted: 10 June 2014 / Published online: 9 July 2014 Ó Springer Science+Business Media Dordrecht 2014 Abstract The Austral papaya (Vasconcellea chilen- molecular markers, along with morphological trait sis) is an endangered species that has valuable analyses, suggest that these relict populations are the characteristics for introgression into other papaya result of a relatively recent fragmentation. This species. These characteristics include disease resis- implies that the fragmentation has not yet had its full tance, cold tolerance and latex with low proteolytic effect on the genetic variation and so emphasises the activity. It is a species that grows under extreme need for clear and urgent conservation measures to environmental conditions of drought, salinity and preserve the remaining genetic variation, particularly temperature; it is found growing naturally in Chile but for the most northern of the three populations which is today only as three remnant populations. The results presently unprotected. presented here, using inter simple sequence repeat B. Carrasco Á C. Dı´az H. Silva Facultad de Agronomı´a e Ingenierı´a Forestal, Pontificia Laboratorio de Geno´mica Funcional y Bioinforma´tica, Universidad Cato´lica de Chile, Vicun˜a Mackenna 4860, Departamento de Produccio´n Agrı´cola, Facultad de Macul, Santiago, Chile Ciencias Agrono´micas, Universidad de Chile, Av. Santa e-mail: [email protected] Rosa 11315, 8820808 La Pintana, Santiago, Chile e-mail: [email protected] R. Garcı´a-Gonza´les (&) Á P. Ca´ceres Facultad de Ciencias Agrarias y Forestales, Centro de P. D. S. Caligari Biotecnologı´a de los Recursos Naturales (CENBio), Centre of Plant Breeding and Phenomics Center, Instituto Universidad Cato´lica del Maule, Av. San Miguel 3605, de Ciencias Biolo´gicas, Universidad de Talca, Talca, Chile 2 Norte 685, Talca, Chile e-mail: [email protected] e-mail: [email protected] P. A´ vila Servicio Agrı´cola y Ganadero, 2 Poniente 1180, Talca, Chile e-mail: [email protected] G. A. Lobos Facultad de Ciencias Agrarias, Plant Breeding and Phenomics Center, Universidad de Talca, 2 Norte 685, Talca, Chile e-mail: [email protected] 123 1424 Genet Resour Crop Evol (2014) 61:1423–1432 Keywords Vasconcellea chilensis Á Austral papaya Á Vasconcellea sp. have a small genome, similar to Palo gordo Á Genetic diversity Á ISSR markers Á Carica papaya (between 442.5 and 625.9 Mb) orga- Genetic structure Á Introgression Á Genetic nized into 18 chromosomes, and a pair of primitive sex fragmentation chromosomes (Gschwend et al. 2013). In Chile only two species are known to exist, the first one is the introduced Vasconcellea pubescens (Lenne´ et C. Koch) Badillo which is cultivated in order Introduction to obtain canned fruit and juices. The second one is the endemic Vasconcellea chilensis (Planch. ex A. DC.) Caricaceae is a tropical and subtropical family Solms (Palo gordo, native papaya, Austral papaya), consisting of six genera (Carica, Vasconcellea, Ja- which is a small shrub 1–2 m tall; with a thick and caratia, Cylicomorpha, Jarilla and Horovitzia) with succulent stem (Fig. 1). Its shiny leaves display a very around 35 species. Most of them are found in the variable shape from an oval and deltoid to heart Americas and only the Cylicomorpha genus is native shaped. It has small purple flowers (5–6 mm) that to Africa (Stevens 2001; Gschwend et al. 2013). appear during winter. As most Caricaeae, V. chilensis According to Antunes-Carvalho and Renner (2012), is a polygamous species with male, female and the family belongs to subtropical and tropical regions hermaphrodite plants. The fruits are small with a of North and South America and Africa. The genus diameter around 1–3 cm; the pulp is minimal and the Cylicomorpha, is located in Central Africa (Antunes- latex has low levels of papain activity; the number of Carvalho and Renner 2012). The genus Carica is seeds per fruit ranges from 5 to 10 (Hechenleitner et al. represented by one species, probably the most culti- 2005). vated species of the family. The genus Jacaratia The natural populations of V. chilensis correspond includes seven tree species distributed in Mexico and to the most southerly distributed papaya so far South America (Antunes-Carvalho and Renner 2012). reported. Currently, the fragmented populations are There are two herbaceous genus, Horovitzia, repre- located between the Atacama (28°390S; 71°420W) and sented by Horovitzia cnidoscoloides (Lorence and Valparaiso (33°090S; 71°420W) Regions (Cuevas R.Torres) V. M. Badillo, and Jarilla which has three 1991; Jordan 2011). But in this range there are (to species (Antunes-Carvalho and Renner 2012). Vas- our knowledge) now only three remnant populations, concellea, or ‘‘highland papaya’’, has the largest with more than thirty plants, which is considered as a number of species (21), including a naturally occur- minimum number of individuals to detect alleles ring hybrid, Vasconcellea 9 heilbornii (V. M. Badillo) which are present at a frequency of 0.05 (Sjo¨rgren and V. M. Badillo (Antunes-Carvalho and Renner 2012). Wyo¨ni 1994; Llewellyn et al. 2003). Any other Vasconcellea species are mostly cultivated at high previous small populations are thought to have altitudes in South America, especially in Ecuador, disappeared due to habitat destruction, goat grazing Colombia, and Peru´ (Scheldeman et al. 2011). and harvest for firewood (Serra et al. 1986; Cuevas Because of the botanical similarity with Carica 1991). papaya L., and because they inhabit zones of high Usually the species is found growing under extreme altitude, they are known as highland papayas (National conditions throughout its natural range, such as living Research Council 1989; Badillo 2000; Van Dro- on soils with high salt concentration and subject to ogenbroeck et al. 2004; Scheldeman et al. 2007). drought. Also, it can support large fluctuations in day Botanically, Vasconcellea species have been and night temperatures. In its natural habitat, it shows described as herbaceous open pollinated species, in a very slow growth rate, scarce biomass production which it is possible to distinguish male, female and and its seeds are difficult to germinate. hermaphrodite plants (Storey 1976;Sa´nchez 1994). In the past the species was used for animal feed, Also they display a short generation time (1 year from which is still a current practice (Rı´os 2004). In the seed to productive tree), a high diversity of growing main populations, goats used to feed from the natural habits, papain production, fruit shape, size and colour, populations during the summer, when other vegetation among other attributes (Antunes-Carvalho and Renner was sparse. However, the continuing use as animal 2012). In genetic terms, it is believed that feed is now very limited because the low number of 123 Genet Resour Crop Evol (2014) 61:1423–1432 1425 Fig. 1 Photographs of Austral papaya (Vasconcellea chilensis): A Isolated tree located on Valle del Encanto; B Fruits from V. pubescens (yellow fruited) and V. chilensis (green–brown fruited); C Vasconcellea chilensis leaf,and D fruit morphological variability individuals in natural population, the low rates of seed abiotic stresses in its natural habitats, means the production and lack of application of any agronomic species could be used as a genetic donor to improve its technology in the species. The fruit of the species is agronomic performance, such as cold tolerance and also considered of interest in terms of its fruit disease resistance (Ocampo et al. 2006; Jordan 2011). properties and papain content because its close At the same time, as with many other papaya relation with tropical papaya and highland papaya species, future industrial uses based on its fruits can be (Jordan 2011). Furthermore, its resistance to the expected if the species can be developed and planted 123 1426 Genet Resour Crop Evol (2014) 61:1423–1432 Fig. 2 Map of sampled 71° Vasconcellea chilensis 30° La Serena populations in the semi-arid Huachalalume regions of Coquimbo, Chile Fray Jorge Ovalle Valle del Encanto 31° on a commercial scale (Rodrı´guez et al. 2005), Materials and methods especially in areas where other crops would not grow readily. Sampling of plant material Currently, V. chilensis is considered as an endangered species being classified as ‘‘vulnerable’’ The three major remnant populations of Vasconcellea (Squeo et al. 2001); but it has not been given any chilensis were sampled over a 600 km2 range between economic importance. However, in reality, its latitudes 30° and 31° South, in the semi-arid regions of capacity for living in extreme environmental condi- Coquimbo Region, Chile (Fig. 2). The number of plants tions and low papain activity make it an important sampled was 20, 30 and 61 individuals respectively in wild resource. the three populations, dictated by the number of plants In this work, inter simple sequence repeats (ISSR) existing in each, as shown in Table 1. For genetic study markers were used to assess the genetic structure of the and morphological characterization, samples within V. chilensis populations. ISSR markers are a very each population were collected at random. useful tool for studies of genetic diversity (Wolfe et al. Young leaves collected from the 111 adult plants 1998; Gilbert et al. 1999). They are based on the use of were selected for DNA extraction. The collected microsatellite sequences as primers to generate multi- samples were frozen in liquid nitrogen and stored at ple band patterns. The necessity for previous knowl- -80 °C until DNA extraction.
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