2.4.1. Apple
.
2.4.1. Apple
The cultivated apple, Malus domestica Borkh., is a diploid or triploid species with a haploid set of 17 chromosomes and a genome size of approximately 600 Mb
[150]. Apple cultivation and production constitute one of the major fruit-producing industries addressing markets worldwide. However, climatic changes introduce a series of environmental stressors which challenge apple yield and fruit quality. Apple breeding could greatly benefit from apple wild relatives to face the challenge from adverse environmental conditions and biotic and abiotic stressors
[149]. For example, wild relatives, Malus floribunda, Malus baccata, and Malus micromalus, have been used to pyramid apple scab and powdery mildew resistance genes into progeny
[151]. Assessment of a broad range of wild Malus germplasm over the last 30 years has revealed ample potential sources of resistance to a multitude of diseases. Similarly, apple wild relatives in Malus collections have been evaluated and shown to possess traits related to fruit quality as well as abiotic stress resilience, such as cold hardiness and drought tolerance
[44]. In addition, investigations on the molecular basis of stress tolerance have indicated a key role for a DREB2 (dehydration-responsive element-binding factor 2) homologue in response to drought, cold, and heat in two highly drought-tolerant wild apple relatives, Malus sieversii and Malus prunifolia
[45][46]. Likewise, Diacylglycerol kinase (DGK) genes were found to exhibit marked upregulation in response to drought and salt stress in M. prunifolia [58]. Comparative analyses between two widely used apple rootstocks (M. sieversii and R3) under water deficit conditions demonstrated M. sieversii is more tolerant to drought. Transcriptomic analysis of root tissue showed differential expression of stress-responsive genes associated with oxidative stress, signaling pathways in hormone biosynthesis, and transcriptional regulation between the two genotypes, suggesting these genes play a crucial role in root processes that provide drought tolerance
[152]. Moreover, the deciphering of the cold-tolerant wild apple Malus baccata genome identified cold-responsive genes (COR) that will be useful in marker-assisted selection in breeding programs
[47]. Collectively, the Malus wild relatives provide an important genetic resource for incorporating resilience in cultivated apple varieties.
2.4.2. Cranberry
2.4.2. Cranberry
Research focusing on wild cranberry is another example of targeted use of genetic variation in perennial wild populations toward the benefit of breeding resilient varieties
[48][153]. Cranberry (Vaccinium macrocarpon Ait.), a fruit crop of high economic value in North America, Northern Europe, and Asia, often encounters a series of abiotic and biotic challenges, such as frost damage, high temperatures, drought, flooding, and fungal diseases, which lead to severe production losses. Recently, a collection of many wild cranberry (Vaccinium macrocarpon Aiton) accessions from the northern U.S. and Canada was assessed through environmental association analysis and revealed genomic regions linked to potential abiotic stress tolerance. One hundred twenty-six significant associations between SNP marker loci (many of which tagged genes with functional annotations) and environmental variables of temperature, precipitation, and soil attributes were uncovered
[154].
2.4.3. Grapevine
2.4.3. Grapevine
Although the Vitis genus is composed of 60 species, the species used predominately for grapevine cultivation is Vitis vinifera L. Nevertheless, wild Vitis relatives exhibit important traits not found in V. vinifera, such as resistance to the devastating ‘Pierce’s disease’ (PD) caused by the bacterium Xyllela fastidiosa. Breeding programs focused on a PD-resistant grapevine wild relative, Vitis arizonica, to generate PD-resistant lines. Over the years, using V. arizonica x V. vinifera crosses, repeated backcrosses with V. vinifera and marker assisted selection (MAS) techniques, breeders managed to develop breeding grapevine lines with PD resistance and 97% V. vinifera ancestry
[155]. Similarly, to confront two major grapevine fungal diseases, downy mildew (Plasmopara viticola) and powdery mildew (Erysiphe necator), the wild relative Muscadinia rotundifolia was utilized. Crosses with Vitis vinifera and subsequent crosses with other Vitis hybrids resulted in progeny containing genes implicated in resistance to both powdery and downy mildew
[156][157]. On the other hand, molecular studies have begun to elucidate the genetic basis of abiotic stress tolerance displayed by wild Vitis relatives. Overexpression of a stress-related gene from the Chinese wild grape Vitis yeshanensis encoding a universal stress protein, VyUSPA3, was shown to confer drought tolerance to transgenic V. vinifera cv. ‘Thompson Seedless’
[49] (
Table 1). In addition, comparative transcriptomic analysis performed between a coastline wild grapevine (Vitis vinifera L. ssp. sylvestris) accession which is tolerant to high-salinity levels and the commercial rootstock, Richter 110, a salt-sensitive cultivar, revealed differential gene expression profiles upon salinity stress [65] (
Table 1). These findings facilitate the investigation of gene pathways that play key roles in survival under stress conditions and highlight the potential of such grapevine wild relatives as breeding material both for scion and rootstock improvement. In view of the gloomy projections of 56 to 73% loss of suitable land for viticulture in major wine-producing regions by 2050
[158][159], studies have been focusing on grapevine wild relatives with resilience to climate risk
[160]. Recently, associations of wild species SNPs (single nucleotide polymorphisms) with bioclimatic variables and putative adaptation to biotic and abiotic stressors have been explored [173]. In addition, by integrating species distribution models, adaptive genetic variation, genomic load and phenotype, Aguirre-Liguori et al.
[161] predicted certain accessions of the wild grapevine species, Vitis mustangensis, are well-suited for future climates and can contribute to grapevine bioclimatic adaptation. Importantly, commercial rootstocks currently used globally for grapevine grafting were derived from North American wild Vitis species. These rootstocks have been used since the second half of the nineteenth century to save European grapevines from the plague of the soil-borne aphid, phylloxera (Daktulosphaira vitifoliae)
[162]. Moreover, depending on the rootstock, they confer drought, cold tolerance, and disease resistance to grafted grapevine
[163]. Likewise, rootstocks have been used widely for improving other cultivated woody perennials (apple, pear, peach, mango, citrus, etc.). However, in general, relatively few rootstock genotypes are employed in grafting of woody perennial crops. Wild relatives could serve as a significant allele pool for developing new rootstock varieties with advantageous traits that would impart the grafted plant with resilience to environmental stressors
[148].