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  • Author or Editor: Gale McGranahan x
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Luther Burbank began making controlled crosses between walnut species in the late 19th century after hearing about a “supposed natural European hybrid walnut.” He crossed Juglans hindsii (northern California black walnut) × J. regia (Persian walnut) and produced progeny that he named ‘Paradox’ because of its extremely fast growth and other “anomalies.” He also crossed two American species, J. hindsii × J. nigra (eastern black walnut), producing ‘Royal’ walnut progeny that were fast-growing and prolific nut producers. A third interspecific hybrid was a cross between J. ailantifolia (Japanese walnut) × J. regia that resulted in extremely vigorous progeny but was not named. He observed segregation in the F2 populations and described giants and dwarfs as reversions to ancestral forms. Luther Burbank also made selections for walnut scion cultivars and was especially interested in thin-shelled nuts. He collected seeds from a J. regia growing in San Francisco because it produced regularly and had very high-quality nuts with relatively thin but poorly sealed shells. He selected one of its seedlings as ‘Santa Rosa Soft-Shell’ and described it as bearing large crops of nuts that were nearly white with thin shells and delicious white meat. Burbank’s contributions to the walnut industry endure to this day, especially through the widespread use of seedling and clonal ‘Paradox’ walnut rootstocks.

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Blackline disease of walnut, caused by the Cherry Leafroll virus (CLRV), causes fatal necrosis at the union between English walnut (Juglans regia) scions and hypersensitive rootstocks. `Paradox' hybrids between English walnut and Northern California black walnut (J. hindsii) are hypersensitive to CLRV. In 1983, `Paradox' trees were backcrossed to J. regia and J. hindsii. Patch graft inoculations of 111 backcrosses, the parent species and `Paradox' hybrids indicated that hypersensitivity to CLRV is inherited as a single dominant gene. Since breeding efficiency can be improved by the elimination of susceptible seedlings, we began a search for genetic markers tightly linked to CLRV resistance. DNA was sampled from the test population and from germplasm of both J. regia and J. hindsii. These samples were screened using randomly amplified polymorphic DNA (RAPD) markers and bulk segregant analysis. Over 360 random primers have been screened and two loci that are less than 10 map units from the hypersensitivity locus in this population identified. Efforts to find more closely linked markers that flank the hypersensitivity locus are continuing.

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In vitro rooting of three commercial cultivars of Persian walnut (Juglans regia L.), `Sunland', `Chandler', and `Vina', was examined using a two-phase rooting procedure: root induction in the dark on Murashige and Skoog (MS) medium with 15 μm IBA followed by root development in the light on a mixture of one-quarter strength Driver Kuniyuki Walnut (DKW) basal medium and vermiculite (1:1.25, v/v). Rooting percentages were: `Sunland' (94%), `Chandler (55%), and `Vina' (27%). A positive relationship was observed between the vigor of cultivars and rooting ability, but shoot length did not affect rooting success. Rooting was optimum when shoots were cultured on root induction media for 6 to 8 days. Increasing the sucrose level in the root induction medium to 40 g·L-1 improved rooting, and shoots induced to root at 22 °C rooted more readily than those induced at 30 °C. Either increasing or decreasing the nitrogen level in the multiplication medium had a negative effect on rooting. Rooted walnut shoots often cease growth during acclimatization, resulting in shoot rosetting. Spray application of Promalin® (25 mL·L-1) caused buds to break and induced elongation of shoots. Chemical name used: indole-3-butyric acid (IBA).

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We characterized a population of hybrids between English walnut and Northern California black walnut (Juglans regia X J. hindsii) and their backcrosses (BC) using both genomic markers and morphological traits. ANOVA and regression methods were used on three years' data to identify a subset of five variables that describe the morphological variability among backcross populations and their parents (R2 = 0.89). Genomic markers were identified using randomly amplified polymorphic DNA (RAPD). A subset of 60 markers specific to the donor species (J. hindsii) were scored in 50 backcrosses to estimate the percent recipient genome in each evaluated BC. The backcrosses were ranked using each method of evaluation; correlation between the ranks was 0.423 and highly significant. Each evaluation method has advantages but neither was able to reliably identify elite progeny.

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To determine if flavor differences could be detected among several Persian walnut (Juglans regia L.) cultivars, difference tests with eight cultivars were conducted using the duo-trio method. No differences were found when `Hartley' was compared to `Vina', `Scharsch Franquette', and `Mayette'. However, `Chandler', `Chico', `Howard', and `Sunland' were significantly different, and paired comparisons were then used to test these cultivars against `Hartley' in terms of several flavor characteristics. No differences in astringency and “walnut flavor” were detected; however, `Chandler' was judged to be sweeter than `Hartley', which was sweeter than `Howard'. `Chico' was found to be the firmest cultivar.

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RFLP markers were used to study genetic diversity among California walnut (Juglans regia L.) cultivars and germplasm collected worldwide. 16 of 21 RFLP markers were polymorphic in the 48 walnut accessions tested. Seven RFLP markers permitted unique identification of all walnut cultivars. All genotypes were heterozygous at approximately 20% of the loci for both California and worldwide germplasm. California walnut germplasm contained 65% of the worldwide allelic diversity. Cluster analysis of genetic distance between accessions and principal component analysis of allelic genotypes showed two major groups of walnut domestication. California germplasm was associated with germplasm from France, Central Europe, and Iran, and had less genotypic similarity with germplasm from Nepal, China, Korea, and Japan.

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Abstract

In vitro propagation of mature cultivars of Juglans regia L. (Persian or English walnut) could have several major advantages over propagation by grafting on seedling black walnut [J. hindsii (Jeps.) Jeps.] rootstock. If commercial cultivars perform satisfactorily on their own roots, the expensive and time-consuming process of grafting could be avoided, genetic uniformity could be assured, and trees would not be subject to the lethal girdling of blackline disease (5). In addition, mature clones on their own roots might be more precocious than grafted seedlings. These factors are becoming increasingly important as walnut growers change to high-density or hedgerow orchards (D.E. Ramos, personal communication).

Open Access