RFLPs were studied in 41 populations of 13 Juglans species to study genetic diversity and phylogenetic relationships. 19 single locus nuclear RFLP loci were used to generate genetic distance/identity matrices based on allele frequencies. 21 probes were used to generate genetic distances and phylograms using shared-fragments with parsimony analysis. Parsimony analysis on fragment data produced a minimal length tree in general agreement with distance data trees, but with additional phylogenetic resolution resembling previous systematic studies. All analyses indicate an ancient origin of J. regia, which has been considered a recently derived species. A 10x difference in heterozygosity was seen among species. Genetic differentiation among conspecific east Asian populations was larger than among east Asian species. The opposite was true for American species. J. hindsii is classified as a distinct species and J. cinerea was included in section Cardiocaryon rather than Trachycaryon, from the diversity analysis.
Robert G. Fjellstrom and Dan E. Parfitt
Robert G. Fjellstrom and Dan E. Parfitt
32 cloned probes from a walnut (Juglans sp.) PstI random genomic library were used to develop a linkage map for walnut. Low copy number walnut random genomic DNA probes were constructed and hybridized to restriction endonuclease digested DNA from parent walnut trees from a backcross of (J. hindsii × J. regia) with J. regia to identify parental polymorphism. 63 backcross progeny were analyzed to determine the inheritance and linkage of 48 RFLP loci. 66% of the probes detected duplicated, but unlinked loci. 42 of the RFLP loci could be placed on 12 linkage groups. The other 6 loci could not be placed on common linkage groups. (Theoretical maximum number of linkage groups is 16.) A Poisson probability method for estimating genome size was utilized to calculate the approximate walnut genome length as 1660 cm and to estimate that 138 markers would be needed to cover 95% the walnut genome within 20 cM of each marker.
John P. Edstrom*, William Krueger, and Wilbur Reil
Orchard hedgerow production systems have been used successfully in fruit and nut crops in California for decades to enhance yield, particularly in the early years of production. English walnuts (Juglans regia) are compatible with hedgerow techniques under prime soil conditions but are thought to require deep well drained soil to be commercially productive. Combining the production techniques of micro-irrigation, close spacing, minimal pruning and frequent fertilization in almonds has improved yield substantially on soils exhibiting a shallow, course textured topsoil underlain with a dense clay layer. Paradox hybrid rootstock (J. regia × J. hindsii) has shown greater tolerance to root lesion nematodes and heavier textured or poorly drained soils than Northern California Black (J. hindsii). Fourteen years of evaluation (1986-99) using `Chandler' and `Howard' Ctvs English walnuts in a replicated field trial on marginal soil has shown that 1) yields of 6700 kg·ha-1 (inshell) are attainable under these substandard soil conditions 2) Paradox hybrid rootstock out-yields Northern California Black by 30% on both cultivars tested, 3) kernels of high commercial quality for can be produced for both cultivars and 4) slip plow soil modifications may not improve tree growth, yield or crop quality in drip irrigated walnut hedgerow plantings.
Richard P. Buchner*, Allan Fulton, Bruce Lampinen, Ken Shackel, Terry Prichard, Larry Schwankl, Sam Metcalf, and Cayle Little
Ninth leaf California Chandler Walnuts (Juglans regia) on Northern California Black (Juglans hindsii) or Paradox (English/black hybrid) rootstock were irrigated to achieve three levels of Midday Stem Water Potential (MSWP). Target potentials were: 1) low water stress (average MSWP of -3.2 bars); 2) mild water stress (average MSWP of -6.2 bars); and 3) moderate water stress (average MSWP of -7.3 bars). Stem Water Potential was measured midday (12-4 pm) by placing leaves inside water impervious, light blocking foil bags. Leaves remained bagged for at least ten minutes to achieve equilibrium. Bagged leaves were removed, placed inside a pressure chamber and stem water potential was measured at endpoint. Data are presented for the 2002 and 2003 seasons. Withholding irrigation water had a significant impact on `Chandler' growth, productivity, and profitability particularly on young, vigorously growing trees. Chandler/Black appears to be more tolerant to water stress compared to Chandler/Paradox For Chandler on Paradox, water stress significantly reduced growth, yield, price per pound, percent edible kernel, and resulted in darker kernels. In addition, water stress significantly increased the total percent offgrade. Withholding irrigation does not appear to be a good strategy in young, vigorously growing `Chandler' orchards. Mature trees and trees grafted onto Northern California black rootstock may be more tolerant of moisture stress.
Xiaobo Song, Shengke Xi, Junpei Zhang, Qingguo Ma, Ye Zhou, Dong Pei, Huzhi Xu, and Jianwu Zhang
excellent walnut rootstocks for walnut industry growing in China. Origin ‘Zhong Ning Sheng’ (ZNS) is the result of a cross between JH-8202 ( Juglans hindsii ) and LNJR-01 ( Juglans regia ) performed at a seedling field in Luoning County, Henan, China. The
Gerald S. Dangl, Keith Woeste, Mallikarjuna K. Aradhya, Anne Koehmstedt, Chuck Simon, Daniel Potter, Charles A. Leslie, and Gale McGranahan
One hundred and forty-seven primer pairs originally designed to amplify microsatellites, also known as simple sequence repeats (SSR), in black walnut (Juglans nigra L.) were screened for utility in persian walnut (J. regia L.). Based on scorability and number of informative polymorphisms, the best 14 loci were selected to analyze a diverse group of 47 persian walnut accessions and one J. hindsii (Jepson) Jepson ex R.E. Sm × J. regia hybrid (Paradox) rootstock. Among the 48 accessions, there were 44 unique multi-locus profiles; the accessions with identical profiles appeared to be synonyms. The pairwise genetic distance based on proportion of shared alleles was calculated for all accessions and a UPGMA (unweighted pair group method with arithmetic mean) dendrogram constructed. The results agree well with what is known about the pedigree and/or origins of the genotypes. The SSR markers distinguished pairs of closely related cultivars and should be able to uniquely characterize all walnut cultivars with the exception of budsports. They provide a more powerful and reliable system for the molecular characterization of walnut germplasm than those previously tested. These markers have numerous applications for the walnut industry, including cultivar identification, verification of pedigrees for cultivar and rootstock breeding programs, paternity analysis, and understanding the genetic diversity of germplasm collections.
R. Fjellstrom and D. E. Parfitt
RFLP analysis was employed to study the inheritance of and genetic diversity identified by cloned walnut genomic probes. An interspecific backcross population of (J. hindsii × J. regia) × J. regia was used to determine the inheritance of thirty low copy number RFLP cloned probes. Of these probes, approximately 20% correspond to single copy loci, 40% correspond to single major loci with multiple minor loci, and 40% correspond to two major loci. Twenty of these probes were used to analyze variability within and between 13 walnut species (Juglans spp.). Substantial genetic variation was identified within many wild walnut species, while limited variation was identified within butternut (J. cinerea) and the widely cultivated English walnut (J. regia). Extensive polymorphism was found between walnut species, allowing a phylogenetic relationship of walnuts based upon RFLP markers to be developed. Identification of clonally propagated walnut cultivars by RFLP typing was readily performed in black walnut (J. nigra) accessions, was more difficult in English walnut accessions, and rarely possible in butternut accessions.
J.R. McKenna and E.G. Sutter
Experiments with field-grown hybrid Paradox (Juglans hindsii × J. Regia) walnut trees were conducted to assess the effects of stock plant water status, auxin application method, and the addition of spermine on adventitious root formation in stem cuttings. A 2-fold increase in rooting was noted when semihardwood cuttings were collected from dry (midday Ψw = –1.3 MPa) stock plants compared to the same trees six days later when fully hydrated (midday Ψw = –0.6 MPa). Spermine, when combined with potassium indolebutyric acid (KIBA) and applied as a quick dip, enhanced the rooting percentage in hardwood cuttings (54%) compared to controls treated with KIBA alone (18%). Spermine had no effect when it was applied together with KIBA using a toothpick application, producing 65% rooting compared to controls which had 75% rooting. By itself, spermine had no effect on rooting. The toothpick method for applying rooting compounds resulted in significantly higher rooting percentages for hardwood cuttings, but not for semihardwood cuttings. Combining spermine with KIBA had no effect on rooting of semihardwood cuttings.
William H. Olson and David Ramos
The Persian or English walnut (Juglans regia) is widely cultivated, with commercial production in France, Italy, Turkey, China, and the United States. Practically all of the U.S. production of Persian walnuts is in the central valley of California, which now has about 169,000 bearing acres with an average yield of around one and one-third short tons per acre. Many orchards produce over two tons, and three tons per acre are common in many modern plantings. Walnuts have two major outlets: the exported in-shell market (about 35% of production) and the domestic shelled market (about 68% of production). A cooperative handles about half the crop, while several independent handlers sell the remainder. Walnuts are sensitive to both low and high temperatures. Temperatures in excess of 90 °F will begin to sunburn nuts. Freezing temperatures will damage tender growth in the spring and fall. Dormant trees can tolerate 15 °F without injury if soils are moist. Dry winter soils and cold temperatures cause winter kill. A minimum of 800 hours of winter chilling are required to avoid delayed bud break and poor crops. Walnuts do best on deep, medium textured, well drained soil. Under these conditions, both rootstocks, the Northern California Black Walnut (J. hindsii) and Paradox (J. regia x J. hindsii), do well. Under less favorable soil conditions, Paradox is the preferred rootstock. A mature walnut orchard requires 4 to 4.5 acre-feet of water per acre per year if the trees are to produce the maximum number of high quality nuts possible. Hartley, preferred for its in-shell quality, is the leading cultivar, with about 30% of the acreage. In recent years, the Chandler variety has accounted for most new plantings. It is known for high kernel quality and yields. Yield factors include: bearing habit, bearing area, flower differentiation, fruit set, nut size, kernel percentage, and kernel quality. Major insect pests of walnut include codling moth, navel orangeworm, and walnut husk fly. The major diseases are walnut blight, deep bark canker, Phytophthora, and blackline. Major research efforts include the walnut breeding program, which includes blackline and Phytophthora susceptibility of new cultivars and root-stocks, codling moth and walnut husk fly control, epidemiology and control of walnut blight, pruning and planting strategies, and clonal propagation.
Janine K. Hasey, Dave Ramos, Warren Micke, and Jim Yeager
In a comparison of six walnut rootstocks either nursery-grafted or field-grafted to `Chandler' (Juglans regia), the highest-yielding trees after 9 years are on either seedling or clonal Paradox rootstocks. Trees growing on both Paradox rootstocks had higher yield efficiency than trees on the black rootstocks in both 1995 and 1996. Since 1993, relative tree size based on trunk circumference has not changed: southern California black (J. californica), seedling Paradox and northern California black (J. Hindsii) have remained significantly larger than clonal Paradox, Texas (J. microcarpa) or Arizona (J. major) black rootstocks. The smaller size of clonal as compared with seedling Paradox trees might be explained by a delay in field grafting success. Although both northern and southern California black rootstock trees were significantly larger than clonal Paradox trees, they did not differ significantly in yield and had significantly lower yield efficiency in 1996. Clonal Paradox trees have significantly smaller nut size than northern California black rootstock trees that can be explained by its higher yield efficiency. An adjacent trial planted in 1991 compares micropropagated `Chandler' on its own root vs. `Chandler' on seedling Paradox rootstock. In 1995 and 1996, own-rooted `Chandler' had significantly greater trunk circumference, yield, and yield efficiency than did `Chandler' on Paradox rootstock. Many of the trees on Paradox rootstock are growing very poorly compared to the own rooted trees. This could be due to diversity within the Paradox seed source. If own-rooted `Chandler' trees become commercially available, they may have potential in areas where other rootstocks are undesirable because of hypersensitivity to cherry leafroll virus.