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And Intraspecific Hybrids of Viola 3Wittrockiana and Viola Cornuta

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And Intraspecific Hybrids of Viola 3Wittrockiana and Viola Cornuta HORTSCIENCE 53(9):1300–1305. 2018. https://doi.org/10.21273/HORTSCI13098-18 from the crossing of Viola L. sect. Melanium Ging. species (pansies), V. tricolor L., V. lutea Hudson, and V. altaica Ker-Gawler Karyologic and Heterosis Studies of the (Kroon, 1972). The species differs from wild pansies based on the absence of dark lines on Artificial Inter- and Intraspecific the flower petals (nectar guides for pollina- tors) and the presence of huge blocks of color Hybrids of Viola 3wittrockiana and on the lower (anterior) and lateral petals called a ‘‘face.’’ This pansy and its progeny Viola cornuta have become popular with gardeners and breeders throughout Europe. At the turn of Xiaohua Du1 and Mengye Wang the 20th century, Charles Stewart bred clear Henan Institute of Science and Technology, Xinxiang, Henan Province, color pansies without faces that were wel- comed by North American gardeners. Cur- 453003, China rently, V. ·wittrockiana is one of the Aneta Słomka dominant hybrids among ornamental bedding plants in the U.S. market (Janick, 1998). Department of Plant Cytology and Embryology, Jagiellonian University, 9 During the past 60 years, much of the Gronostajowa St., 30-387 Cracow, Poland innovative pansy breeding has been per- formed in Germany, the United States, and Huichao Liu Japan (Zhang et al., 2010). New pansy Henan Institute of Science and Technology, Xinxiang, Henan Province, varieties with different colors, such as shades 453003, China of pink, rose, or orange, and unusual bicolor designs have been introduced and are avail- Additional index words. pansies, inbred lines, aneuploid chromosome numbers, artificial able to gardeners. Studies using a simple crossing, flower size increase, F1 progeny selection method to reduce the ploidy level Abstract. An artificial crossing was made between four Viola 3wittrockiana (2n = 48) lines by crossing with diploids to improve unifor- and one Viola cornuta inbred (2n = 26) line as well as among V. 3wittrockiana lines to mity, as well as an increase of polyploidy by investigate the cytologic characteristics, fertility, and heteroses of hybrids. The sizes of mutagenesis to maximize plant morphologic flowers and stomata and the leaf areas of the V. 3wittrockiana plants were larger than and physiologic characters, have already those of the V. cornuta plants. The karyologic stability of V. cornuta was greater than that been performed (Dalbato et al., 2013; Kroon, of V. 3wittrockiana, and ’20% of individuals in the latter deviated from the given 2n = 1972; Lagibo et al., 2005). Lagibo et al. (2005) demonstrated the positive influence 48. Capsule setting after interspecies hand pollination ranged from 28% to 94%, and F1 seed germination ranged from 16% to 88%, revealing no obvious pre- or postzygotic of increased polyploid level on frost toler- ance and winter survival of garden pansy. selection mechanisms. The chromosome numbers in V. 3wittrockiana 3 V. cornuta F1 seedlings varied (2n = 34–38), with about half the cells exhibiting the intermediate (2n = Along with the increasing popularity of 36) value. These interspecific hybrids exhibited positive heterosis in flower size (range, pansy cultivation worldwide, breeding goals 11% to 66%) and negative heterosis in plant height (range, –12% to –57%). Both of these continue to develop. Recently, V. cornuta L. traits are desirable in pansy breeding programs. (horned pansy) has been crossed successfully with V. ·wittrockiana for the creation of new desirable phenotypic traits, such as increased flowering, plant compactness, and tolerance Breeding methods for ornamental plants are F1 hybrids produced by hand crossing. range from traditional selection breeding Currently, such production of Viola tricolor to environmental stresses (Li, 2010; Liu, programs to modern biotechnological and maxima with hermaphrodite flowers requir- 2008; Zhang and Gu, 2009). Currently, some genetic engineering methods. Many orna- ing emasculation occasionally occurs at the hybrid cultivars of these two species—e.g., mental plants, including geraniums, impa- expense of limited profitability (De, 2017; ‘Nature Series’ and ‘Power Mini’—are bred tiens, begonias, and primulas, originated Reimann-Philipp, 1983). by the Japanese seed company Takii Seed from interspecific and intergeneric crosses, Heterosis is a phenomenon whereby the (Kyoto, Japan) and Sakata Seed (Japan) re- which leads to a high degree of heterozy- phenotype of F hybrids is superior to that of spectively. To meet the ever-growing expec- 1 tations of customers (practical approach) and gosity in the resulting hybrids, with polyploidy their parents. Intralocus interactions between to increase the predictability of the crossing and aneuploidy often occurring. In seed- alleles, complementation of dominant alleles, treatments (basic science approach), we propagated ornamentals with a considerable or interloci epistatic interactions are genetic adopted four inbred lines of V. ·wittrockiana seed market, cultivars will normally be bred mechanisms that may cause nonadditive and one inbred line of V. cornuta to in- as hybrids based on inbred parental lines. The phenotypic inheritance in hybrids. However, main emphasis in ornamental plant breeding vestigate the cytologic basis of hybridization different views exist on what portion of the and heterosis in their hybrids. is to improve the variety traits, novel color, heterotic variation is modulated by each of form, size, number of flowers, flower vase these mechanisms (Fridman, 2015). Materials and Methods life, repeat blooming, disease resistance, Viola ·wittrockiana Gams ex Kappert., nutrient uptake capacity, and growth habits known as garden pansy, is one of the most · (De, 2017). Commercially available pansies Derivation of inbred lines of V. wittrockiana common bedding plants worldwide, exhibit- and V. cornuta and their morpho-anatomic ing rich flower colors, cold tolerance, and description. The breeding experiments were other charming characteristics, such as odor performed in the garden field of the Henan Received for publication 2 Apr. 2018. Accepted for and taste (Fernandes et al., 2017; Lagibo Institute of Science and Technology (Xin- publication 20 July 2018. et al., 2005; Li et al., 2014, and literature xiang, China) from 2015 to 2017. Four V. This research was supported by the Henan Province cited therein). This species is also studied as ·wittrockiana inbred lines and one V. cor- Science and Technology Research Project (no. a promising source of natural antioxidants 152102110088) and the Henan Institute of Science nuta inbred line (Table 1; Fig. 1A–E) were and Technology Provincial and Ministerial Achieve- (Vukics et al., 2008). Present-day garden generated from commercially available seeds ment Award Cultivation Project (2017CG02). pansies are horticultural hybrids rather than by self-crossing for five or six generations, 1Corresponding author. E-mail: duxiaohua0124@ a natural species. Initially, the species was given accession numbers, and used as paren- sina.com. selected as an interspecific hybrid resulting tal lines. Plants were grown in a closed plastic 1300 HORTSCIENCE VOL. 53(9) SEPTEMBER 2018 Table 1. The pedigrees, accession numbers. and origins of Viola cornuta and Viola ·wittrockiana lines further growth and blossoming (Fig. 1F). Ten used for inter- and intraspecific crosses. randomly selected plants of each accession No. Name Pedigree Species Origin were chosen in Apr. 2017 to investigate plant 1 JB-1-1-2 Inbred line derived V. cornuta Goldsmith seed height, plant spread (width), flower number from Penny Blue and size (height), flower shape (height–width 2 DFM-11-2-1 Inbred line derived from V. ·wittrockiana Gartenland GmbH ratio), leaf shape (length–width ratio), and Fruhbl€ uhende€ Mischung Aschersleben area. A fully developed leaf and an open · 3 DFM-11-1-3 Inbred line derived from V. wittrockiana Gartenland GmbH flower were measured per plant. Mid-parent Fruhbl€ uhende€ Mischung Aschersleben · heterosis of these characters was calculated 4 DFM-8-4-3 Inbred line derived from V. wittrockiana Gartenland GmbH · Fruhbl€ uhende€ Mischung Aschersleben according to the Eq. 100 [F1–(P1 +P2)/2]/ 5 XXL-YB-1-1-1-1 Inbred line derived from V. ·wittrockiana PanAmerican seed (P1 +P2)/2 (Marame et al., 2009). extra flower size Chromosome counting in inbred lines and their F1 hybrid progeny. The seeds of inbred lines and their F1 progeny were germinated on filter paper soaked with distilled water at room temperature. Roots 1 to 2 cm in length were sampled, and their root tips were dis- sected under a stereomicroscope. The seed- lings were pretreated for 24 h at 0 °C and fixed in a mixture of glacial acetic acid and absolute ethanol (3:1 ratio, v/v) for 24 h. The fixed root tips were then macerated with 1mol·L–1 hydrochloric acid at 60 °Cfor6to 8 min. Finally, the material was stained with carbol fuchsin, flattened with a cover slip, and observed under a Nikon Eclipse 80i microscope (Nikon Corporation) with a mag- nification of 10 (eye lens) · 100 (field lens). The images were generated using CellSens (ver. 1.12) Imaging Software (Olympus Cor- poration, Tokyo, Japan). At least 30 plates from 10 plants per accession were observed. Fig. 1. (A) Plants of inbred lines of Viola cornuta during flowering. (B–E) Plants of inbred lines of Viola Results ·wittrockiana during flowering. (F) Plants of hybrid of V. cornuta and V. ·wittrockiana during flowering. (A) JB-1-1-2, (B) DFM-11-2-1, (C) DFM-11-1-3, (D) DFM-8-4-3, (E) XXL-YB-1-1-1-1, Morpho-anatomic and cytologic charac- · and (F) XXL-YB-1-1-1-1 DFM-8-4-3. teristics of the inbred lines of V. cornuta and V. ·wittrockiana. Stomata length, stomata width, leaf area, and flower size of the V. chamber from Oct. 2015 to Mar. 2016 and acid paper bags and labeled immediately. cornuta inbred line (118.1 mm, 70.0 mm, 2.5 then under insect netting from Apr. to June One week later, the isolation bags were cm2, and 2.1 cm, respectively) were reduced 2016 to avoid high temperatures and insect removed from the enlarged ovaries, and significantly compared with those of the V.
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