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Ploidy, Relative Genome Size, and Inheritance of Spotted Foliage In HORTSCIENCE 53(9):1271–1274. 2018. https://doi.org/10.21273/HORTSCI13221-18 eriobotryifolia, himalaica,andomeiensis, have been reported to be diploid with 2n = 2x = 16 (Hara, 1966; Kurosawa, 1971; Ploidy, Relative Genome Size, and 1981; The Chromosome Counts Database, 2018). Aucuba japonica aremorevariable Inheritance of Spotted Foliage in and well-studied, with three varieties (ja- ponica, borealis,andovoidea)thathave Aucuba Species (Garryaceae) been classified based on a combination of morphology, ploidy, and geographic dis- 1,5 2 2 Thomas G. Ranney , Tracy H. Thomasson , Kristin Neill , tribution (Ohi et al., 2003). Variety bor- Nathan P. Lynch3, and Mark Weathington4 ealis is the most northerly distributed and Mountain Crop Improvement Lab, Department of Horticultural Science, found in southwestern Hokkaido and the Mountain Horticultural Crops Research and Extension Center, North SeaofJapan(Western)sideofHonshuand is tetraploid (2n =4x = 32). Variety Carolina State University, 455 Research Drive, Mills River, NC 28759- japonica is present on the Pacific Ocean 3423 (Eastern) side of Honshu and Shikoku and Additional index words. cytotype, DNA content, plant breeding, polyploidy is also tetraploid. Variety ovoidea is dis- tributed further south, ranging from south- Abstract. Aucuba have been cultivated for centuries and are valued as adaptable, broad- ern Honshu, western Shikoku, and Kyushu leaved, evergreen shrubs that also can have attractive, spotted variegations on the foliage. to Okinawa, and is diploid. Variety bor- Improved understanding of the cytogenetics and heritability of specific traits, for specific ealis is considered to be the most cold- clones and cultivars, can provide basic information to help facilitate the breeding and hardy of the group with a smaller stature, improvement of aucuba. The objectives of this study were to determine ploidy level and decumbent shoots, smaller leaves, and relative genome size of a diverse collection of species and cultivars of aucuba using flow pubescent young buds and inflorescences. cytometry and cytology and to make additional observations on heritability of spotted Varieties japonica and ovoidea are larger leaf variegation. Chromosome counts were 2n =2x = 16 for Aucuba chinensis (A. growing and more similar to one another omeiensis), 2n =4x = 32 for A. japonica ‘Rozannie’, and 2n =6x = 48 for A. sp. ‘Hosoba’. in appearance but clearly differentiated Relative 2C genome size for the 57 taxa varied from 13.8 pg for A. obcordata to 42.0 pg for by ploidy. Karyomorpholological studies A. ‘Hosoba’ and fell within three discrete groups consistent with cytotype. Genome size on a diverse sampling of diploid A. japonica for diploid taxa (A. chinensis and A. obcordata) ranged from 13.8 to 21.0 pg, tetraploids var. ovoidea showed the presence of XY sex (A. himalaica var. oblanceolata, A. japonica, and A. japonica var. borealis) ranged from chromosomes where the fourth-longest pair 28.8 to 31.2 pg, and the first-ever reported hexaploids (A. ‘Hosoba’ and A. sp. – Vietnam) of chromosomes was heterogeneous in ranged from 40.5 to 42.0 pg. Unlike prior reports that indicated inheritance of spotted male individuals and homogeneous in fe- variegations were extranuclear genes that were maternally inherited, we found that the male individuals (Tsusaka et al., 2007). A. japonica spotted leaf trait expressed in ‘Shilpot’ appears to be a nuclear gene that is Despite the work that has been completed inherited in a quantitative fashion and not strictly maternal. These data provide an on chromosome numbers and ploidy in enhanced foundation for breeding improved aucuba. aucuba, there is extremely little documen- tation of ploidy for specific cultivars and clones. Aucuba (Aucuba) is a relatively small much broader past distribution (Wehr and Minimal work has been published on genus of about nine species (The Plant List, Hopkins, 1994). genetics and heritability of specific traits in 2018). Formerly placed in the Cornaceae Cultivated for centuries, aucuba are re- aucuba. Hagedoorn (1950) first reported that and Aucubaceae families, the genus now vered as showy, adaptable, evergreen shrubs the spotted leaf trait was maternally inherited resides in the Garryaceae (Stevens, 2018). (Creech, 1984). Aucuba japonica is com- and that all seedlings from spotted-leaved Although currently extant in Eastern Asia monly grown around shrines and temples in females had spotted leaves regardless of the (ranging from the Eastern Himalayas east Japan, thriving in the understory of Crypto- male parent and that seedlings from green- to China, Korea, and Japan and south to meria japonica. John Graeffer is credited Myanmar, Taiwan, and Vietnam), fossil leaved female parents always had green with introducing A. japonica to the Western leaves even when the pollen parent had leaves of aucuba from the Eocene have been World in 1783, where it has become an im- found in Washington State, suggesting a spotted leaves. Allen (1990) further substan- portant and valuable landscape plant (The tiated this observation by collecting seed International Dendrology Society, 2018). Of from green-leaved and spotted-leaved plants particular merit are many cultivars that have that all produced seedlings with the same showy, variegated foliage ranging from gold Received for publication 16 May 2018. Accepted phenotype as the female parent. for publication 6 July 2018. flecking and spots to variegated leaf margins. There is considerable potential to breed This work was funded, in part, by the North Although the flowers are inconspicuous and and improve aucuba as nursery crops. The Carolina Agricultural Research Service (NCARS), dioecious, female plants can have outstand- diversity in desirable foliage characteristics Raleigh, NC, the North Carolina Biotechnology ing displays of red fleshy fruit. In addition to (leaf size and variegations), plant size, cold Center, Research Triangle Park, NC, and the Kenan ornamental merit, A. japonica are notoriously hardiness, and potentially disease resistance Institute, Raleigh, NC. tolerant of dry shade, pruning, pollution, and Plant material was graciously provided by the JC and regional adaptability provides substantial Raulston Arboretum, Raleigh, NC, the U.S. Na- general neglect (Lehrer, 2009). Large speci- opportunities to combine desirable traits. tional Arboretum, Washington, DC, and Dan mens frequently are found on old estates and Additional cytogenetic data, with specific Hinkley, Indianola, WA. We would like to thank homesteads in the Southeastern United information on ploidy of specific cultivars the staff at the Mountain Crop Improvement Lab States, where they have long outlived their and clones, and information on the mode of and Mountain Horticultural Crops Research and original gardeners. inheritance of specific traits, would provide Extension Center for their technical assistance. Knowledge of ploidy is important for 1 valuable information to better enable plant JC Raulston Distinguished Professor. plant breeders, as it can influence cross- breeding programs. The objectives of this 2Research intern. 3Research specialist. ability, fertility of progeny, segregation of study were to determine ploidy and rela- 4Director, JC Raulston Arboretum. traits, and gene expression. There is lim- tive genome size for a diverse collection 5Corresponding author. E-mail: tom_ranney@ ited information on the cytogenetics of of species and cultivars of aucuba using ncsu.edu. aucuba. Some species, including A. chinensis, flow cytometry and cytology and to make HORTSCIENCE VOL. 53(9) SEPTEMBER 2018 1271 additional observations on the heritability of Heritability of spotted variegation. Cross spotted leaf variegation of specific cultivars. pollinations were completed between the female, green-leaved cultivar A. japonica Materials and Methods ‘Petite Jade’ and the male, spotted-leaved cultivar A. japonica ‘Shilpot’ in Mar. 2015. Plant material. Plants and/or tissue from Fruit were collected in the Fall of 2015, seeds 57 taxa of aucuba were obtained from the JC were extracted, given cold/moist stratifica- Raulston Arboretum (JCRA), Raleigh, NC; tion for 90 d at 6 °C, and germinated and the Mountain Crop Improvement Laboratory, grown under standard production practices. Mills River, NC; and the United States Seedlings were evaluated for expression of National Arboretum, Washington, DC. Cur- spotted variegations in May 2018. rent nomenclature for aucuba was adopted from The Plant List (2018) and Xiang and Results and Discussion Boufford (2005). Synonyms from accession records and collection data were included The chromosome count for A. chinensis when present. (A. omeiensis, JCRA 120820) was diploid Cytology. Chromosome counts were de- with 2n =2x = 16 (Fig. 1), consistent with termined for three taxa, including A. chinen- a previous report for A. chinensis (Kurosawa, sis (A. omeiensis, JCRA 120820), A. japonica 1981). Aucuba japonica ‘Rozannie’ (MCIL ‘Rozannie’ (MCIL 2015-077), and A. 2015-077) was confirmed to be tetraploid ‘Hosoba’ (JCRA 120812) to determine chro- with 2n =4x = 32 (Fig. 1), consistent with mosome numbers and calibrate genome size other reports of tetraploid forms of A. japon- measurements with ploidy. Actively growing ica var. japonica and var. borealis (Ohi et al., root tips were harvested from container- 2003). Surprisingly, A. ‘Hosoba’ (JCRA grown plants in mid-July and placed into 120812) was found to be hexaploid with a solution of 2 mM 8-hydroxyquinoline and 2n =6x = 48 (Fig. 1), being the first report 0.248 mM
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