Chapter 23 Phalaenopsis

Chia-Chi Hsu, Hong-Hwa Chen, and Wen-Huei Chen

Abstract Phalaenopsis is one of the most popular cultivated orchids worldwide. So far, 92 native species and 34,112 hybrids of Phalaenopsis have been registered in the Royal Horticultural Society (RHS), but only 18 native species are frequently used for breeding. In Phalaenopsis market, large flowers with various colors are most popular. Phal. amabilis and Phal. aphrodite are the major species used for breeding the white-and-large-flower hybrids. The colored hybrids include pink, red-­ to-purple,­ green, yellow-to-orange, and black colors, as well as distinct colors in tepals with red lip. For large-and-red flowers,Phal. schilleriana and Phal. sanderiana are the major parent species, whereas Phal. equestris and Phal. pulcherrima are important for the development of the small- and medium-sized red flowers. Members of the subgenusPolychilos are the most important ancestors for yellow-to-orange flowers. More recently, there is an increased interest in plants with white, pink, or yellow colors with red stripes and/or spots. These traits are introduced from Phal. lindenii, Phal. stuartiana, and Phal. amboinensis. Furthermore, harlequin flowers which have clown-like spots and very complicated color patterns are released in the market, as well as hybrids with peloric and Bigfoot flowers. Besides color, scent and (a)biotic stress resistance are becoming important targets in Phalaenopsis breeding.

Keywords Bigfoot · Breeding · Harlequin · Orchids · Peloric · Phalaenopsis

C.-C. Hsu Department of Life Sciences, National Cheng Kung University, Tainan, Taiwan H.-H. Chen Department of Life Sciences, National Cheng Kung University, Tainan, Taiwan Orchid Research and Development Center, National Cheng Kung University, Tainan, Taiwan e-mail: [email protected] W.-H. Chen (*) Orchid Research and Development Center, National Cheng Kung University, Tainan, Taiwan

© Springer International Publishing AG, part of Springer Nature 2018 567 J. Van Huylenbroeck (ed.), Ornamental Crops, Handbook of Plant Breeding 11, https://doi.org/10.1007/978-3-319-90698-0_23 568 C.-C. Hsu et al.

23.1 Introduction

Phalaenopsis is named with “phalaina” for moth and “opsis” for appearance and is commonly known as “moth orchids”. Phalaenopsis has an unique and beautiful flower morphology. Three sepals are in the outer whorl of the flowers, coming with two petals and one specialized lip. The column with the fused stamen and carpel is located within the center of the flowers. The morphology and colors of sepals and petals are very similar, and together they are called tepals. In contrast, the lip is derived from the specialization of a petal, but its shape and color are very different from petals. The specialized lip has made the flower zygomorphic, and the entire flower is similar to a flying butterfly. Phalaenopsis has an infinite inflorescence, which can have many flowers, and the flowers can last for a long time. All these features made that Phalaenopsis is currently the most successful and popular genus within the Orchidaceae for the ornamental flower market. According to the records in the Royal Horticultural Society (RHS), consulted with the OrchidWiz Orchid Database Software, there are 92 native species of Phalaenopsis worldwide, including the merge of genus Doritis, Hygrochilus, Kingidium, Lesliea, Nothodoritis, Ornithochilus, and Sedirea (OrchidWiz 2017). Based on the characterization of Christenson (2001), these native species can be divided into five subgenera, including Aphyllae, Parishianae, Phalaenopsis, Polychilos, and Proboscidioides. Only 18 species belong to the subgenera Phalaenopsis and Polychilos, but these species are among the most frequently used breeding parents for numerous cultivars. At present, a total of 34,112 hybrids have been bred and officially registered in the RHS database (OrchidWiz 2017).

23.2 Phalaenopsis Markets in the World

Phalaenopsis is the number one potted ornamental plant in the world market (Chen 2017). Based on the survey data from Taiwan Orchid Growers Association in 2014, the orchid growers from the Netherlands and Taiwan nearly dominate the world with their large-scale production and supply of orchid plants.

23.2.1 Phalaenopsis Market in European Union

The European market with 160 million pots sold each year is the biggest in the world. The Netherlands is the major producer of Phalaenopsis orchids in Europe. Different needs can be distinguished for different countries. The Netherlands prefers Phalaenopsis with two or three floral stalks displaying any flower size and color. The consumer in the UK prefers orchids with one long floral stalk showing large flowers (>10 cm in width). People in the Nordic countries adore plants with two 23 Phalaenopsis 569 floral stalks and medium-sized flowers (7~10 cm). The north of Italy prefers the ones with large flowers, while the south Italian people enjoy the ones with small flowers (2~7 cm). Overall for the Phalaenopsis market in Europe, the major trend is to have two to three floral stalks with many flowers in medium size. About 65% of the production are 12 cm pots with 6.5~10 cm flowers, while either 9 cm pots with 6.5~10 cm flowers or 6 cm pots with 4~5.5 cm flowers count for 15%. The large flowers (10~12 cm) occupy around 10% and are produced in 15 cm pots. The costs for plants varied from 1.5~2.5, 3.5, and 4~5 Euro for one, two, and three stalks, respectively. Thus, the hybrids which are prone to produce many floral stalks and flowers are well accepted. Phalaenopsis orchid flowers in various colors are well established in Europe, although the high-intensity and deep colors are preferred. Other important characteristics are easily grown plants, easy flowering, resistance to pathogens, and plants with many stalks and flowers.

23.2.2 Phalaenopsis Market in the USA

In the American market, about 24 million pots are sold each year. The American Orchid Society (AOS) is the biggest association for orchid breeders. Overall, American people prefer large plants with large flowers (8~13 cm) (Table 23.1). People in the east of America love the plants with large flowers and white color, while the western American people prefer the large flowers with various colors. For large flowers, the plants with two floral stalks are the better choices. The inflorescence has to contain more than nine flowers with flower widths more than 11 cm. White flowers occupy 50~70% of the market in the East, including 5% for the white flowers with red lip. The pink and red colors are about 10~20% of the assortment. The ratios of yellow flowers, yellow flowers with red lip, and black flowers have been increased more recently. The medium flowers (5~9 cm in width) have 2 floral stalks with 7 flowers on each stalk and are produced in various colors, including pink, red, purple-to-red,

Table 23.1 The major preferred Phalaenopsis orchids in the USA Pot size Percentage Flower width (cm) Height of floral stalks (cm) East 2″ (6 cm) 5% 4.5~5 25~28 3″ (9 cm) 20% 6~7.5 42~45 4″ (12 cm) 60% 8~10 55~60 5″ (15 cm) 15% 10~13 60~70 West 2″ (6 cm) 5% 4.5~5 25~30 3″ (9 cm) 30% 5.8~7 35~42 4″ (12 cm) 50% 8.5~10 55~60 5″ (15 cm) 15% 10~13 60~70 570 C.-C. Hsu et al.

Table 23.2 Top 10 countries of Taiwan Phalaenopsis plants export in 2014 No. Country Weight (metric ton) Value (thousand US$) Percentage 1 USA 4890 53,943 41.90% 2 Japan 2750 34,284 26.60% 3 Vietnam 1133 6814 5.30% 4 Australia 525 6762 5.30% 5 Netherlands 355 5867 4.60% 6 Canada 482 3734 2.90% 7 England 124 2919 2.30% 8 Korea 152 1913 1.50% 9 La Réunion 175 1678 1.30% 10 Hong Kong 243 1629 1.30% 11 Others 897 9177 7.10% yellow, and orange colors, as well as white flower with red lip or spots, yellow flow- ers with red lip, stripes, or spots. The small flowers are not popular in America.

23.2.3 Phalaenopsis Market in Asia

Totally, the Phalaenopsis market in Asia is estimated at 84 million pots each year, including 60 million in China, 14 million in Japan, and 10 million in Southeast Asia. More than 85% of Phalaenopsis in China are large flowers with red color, while Japanese people prefer the large-and-white flowers (80% of market share). Taiwan is the major exporting country in the region, and also the most new Phalaenopsis hybrids have been bred there (Table 23.2).

23.3 Directions of Phalaenopsis Breeding

23.3.1 Major Ornamental Traits for Phalaenopsis Flowers

Flower color is the most important characteristic. Although white is still dominating the market, various other colors including pink, red, purple-to-red, yellow, orange, green, and black colors are breeding targets. Flowers with various pigmentation patterns, meaning two or more colors present in one flower which forms a pattern, such as the distinct colors in tepals with red lip, or the white or yellow flowers with red stripes or spots, are getting more interest. Based on the flower sizes of the bred hybrids, they can be divided in three groups – large, medium, and small flowers for >10 cm, 7~10 cm, and < 7 cm in width, respectively. Sometimes, the flowers with deranged morphology were found, 23 Phalaenopsis 571 but only the stable and beautiful ones were retained. Otherwise, they were discarded as mutants. Two types of them have been often used, the peloric and the Bigfoot flowers. The peloric flowers contain two petals converted to lip-like petal, while the Bigfoot flowers display a petal-like lip morphology.

23.3.2 Native Phalaenopsis Species for Breeding

Among the 92 native species, only about 18 species are frequently used as breeding parents. They belong to the subgenus Phalaenopsis and the subgenus Polychilos. For the subgenus Phalaenopsis, they comprise Phal. amabilis, Phal. aphrodite, Phal. equestris, Phal. lindenii, Phal. pulcherrima, Phal. sanderiana, Phal. schilleriana, and Phal. stuartiana. For the subgenus Polychilos, they consist from Phal. amboinensis, Phal. bellina, Phal. cornu-cervi, Phal. fasciata, Phal. gigantea, Phal. lueddemanniana, Phal. mannii, Phal. sumatrana, Phal. venosa, and Phal. violacea (Table 23.3). These 18 native species varied in plant sizes, flower morphology, colors, and fragrances. In the subgenus Phalaenopsis, Phal. amabilis and Phal. aphrodite are species with white and medium flowers (7.5~7.8 cm) which are used for breeding the white and large flowers with more than 10 cm in size (Table 23.3). Phal. sanderiana, Phal. schilleriana, and Phal. stuartiana have pink and medium flowers (6.3~7.9 cm), and they are used for breeding the red and large-flower hybrids. In addition, Phal. stuartiana has white flowers with red spots, and it is the major breeding parent for red-spot feature. Phal. equestris, Phal. lindenii, and Phal. pulcherrima have small flowers (2.9~3.6 cm) and various colors. These species are the major breeding parents for small and medium flowers.Phal. equestris provides many flower traits, Phal. lindenii introduces the red-stripe feature, and Phal. pulcherrima gives the dark-red-to-purple colors. In the subgenus Polychilos, species are usually characterized by strong scents as is the case for Phal. amboinensis, Phal. bellina, Phal. lueddemanniana, Phal. venosa, and Phal. violacea. So they are the major parents for breeding the scented hybrids. In addition, Phal. amboinensis, Phal. fasciata, Phal. gigantea, Phal. lueddemanniana, Phal. mannii, Phal. sumatrana, and Phal. venosa have yellow-­ colored flowers, and they are used as parents for breeding yellow flowers. Two varieties of Phal. bellina and Phal. violacea have been found with purple-to-blue color and are used for breeding the dreaming blue hybrids. Except for the differential appearance in floral morphology between the species from both subgenera, also the size of the chromosomes varied. The subgenus Phalaenopsis has small chromosomes, while the subgenus Polychilos has medium to large chromosomes (see Sect. 23.7). The variations in chromosome size hamper successful crossbreeding between the two subgenera due to chromosome pairing problems during meiosis. 572 C.-C. Hsu et al.

Table 23.3 Native Phalaenopsis species used as breeding parents Total no. of Average Average No. of hybrids in hybrids flowers per flower size Species first generation derived Awards spike (cm) Phal. amabilis 475 31,573 94 13.4 7.8 Phal. amboinensis 532 14,951 102 3.6 5.5 Phal. aphrodite 58 31,460 39 16.8 7.6 Phal. 6 6 7 2.7 1.1 appendiculata Phal. bastianii 18 18 13 7.7 3.9 Phal. bellina 104 245 130 2.5 5.1 Phal. braceana 5 5 6 8.5 3.3 Phal. buyssoniana 35 490 15 20.8 4.1 Phal. celebensis 45 65 24 54.8 3.1 Phal. chibae 18 20 11 22.5 1.0 Phal. cochlearis 27 75 3 5.4 4.0 Phal. corningiana 63 215 17 3.4 5.8 Phal. cornu-cervi 99 372 116 4.5 3.0 Phal. deliciosa 31 49 13 18.5 1.6 Phal. doweryensis 5 5 4 5.2 5.2 Phal. equestris 550 21,805 127 28.0 2.9 Phal. fasciata 124 9729 7 3.8 5.0 Phal. fimbriata 56 169 6 4.7 3.8 Phal. finleyi 25 28 5 3.5 1.6 Phal. floresensis 36 77 6 3.3 4.0 Phal. fuscata 103 273 4 13.1 3.8 Phal. gibbosa 4 5 4 3.9 1.1 Phal. gigantea 206 5270 84 22.8 5.4 Phal. hieroglyphica 20 8514 71 4.1 5.6 Phal. honghenensis 22 23 6 12.1 2.9 Phal. hygrochila 43 66 5 5.3 3.5 Phal. 20 21 2 3.1 3.8 inscriptiosinensis Phal. intermedia 6 8 2 10.0 4.0 Phal. javanica 99 187 9 3.4 2.7 Phal. kunstleri 8 9 3 5.9 3.9 Phal. lindenii 115 609 40 22.6 3.6 Phal. lobbii 55 104 64 4.9 2.0 Phal. lowii 10 16 8 7.8 3.9 Phal. 325 20,024 56 3.3 5.5 lueddemanniana Phal. maculata 34 79 8 4.3 3.2 Phal. malipoensis 3 3 2 6.0 1.6 Phal. mannii 157 1212 46 10.7 3.4 (continued) 23 Phalaenopsis 573

Table 23.3 (continued) Total no. of Average Average No. of hybrids in hybrids flowers per flower size Species first generation derived Awards spike (cm) Phal. mariae 107 1224 20 10.2 3.7 Phal. marriottiana 7 7 3 6.8 4.8 Phal. 5 16 1 3.0 4.4 mentawaiensis Phal. micholitzii 50 1211 5 1.6 4.9 Phal. modesta 40 48 7 3.8 3.1 Phal. pallens 25 497 7 2.7 5.1 Phal. pantherina 16 29 3 2.2 4.4 Phal. parishii 31 73 24 5.3 1.6 Phal. 106 162 17 26.5 8.1 philippinensis Phal. pulcherrima 278 9466 143 22.1 3.2 Phal. pulchra 37 47 5 12.7 5.0 Phal. 5 5 2 1.8 4.7 reichenbachiana Phal. sanderiana 114 25,426 6 16.3 7.9 Phal. schilleriana 246 27,991 71 34.7 7.0 Phal. speciosa 26 112 10 2.5 4.8 Phal. stobartiana 10 10 7 31.6 3.1 Phal. stuartiana 392 26,377 42 34.7 6.3 Phal. sumatrana 166 4485 28 4.1 5.8 Phal. taenialis 10 13 6 13.3 3.0 Phal. tetraspis 71 129 31 3.5 4.5 Phal. thailandica 10 12 1 6.3 1.3 Phal. venosa 333 3161 29 3.2 4.2 Phal. violacea 472 5040 197 2.7 5.0 Phal. viridis 14 16 4 4.9 4.1 Phal. wilsonii 33 33 8 8.8 3.0 Data were collected from OrchidWiz X3.3, 2017

23.3.3 Breeding Directions for Phalaenopsis

The breeding directions can be roughly separated into three major features: flower morphology, various flower colors, and scent. In addition, selection toward plants with increased resistance to pathogens and extreme environments is performed in the bred hybrids, but so far it is not the major purpose of most breeding programs. In flower morphology, large flowers are the major breeding direction, while a new trend toward small hybrids has also been initiated. Breeding for flowers with changed morphology such as the peloric flowers with petal becoming lip-like shape or the Bigfoot with the lip turning into petal-like morphology is started recently. 574 C.-C. Hsu et al.

For flower colors, the breeding for white, pink, and red hybrids is almost satu- rated, and now the orchid breeders have turned to find the flowers with various pigmentation patterning, such as red spots and stripes distributed in different regions. Rare and new colors are also preferred, such as yellow, orange, green, and black, as well as the dreaming true-blue color, but not the purple-to-blue color. So far, the true-blue color is still not yet found in both native and bred Phalaenopsis orchids. Due to unique and spectacular floral morphology as well as abundant variations in flower color, the trait of scent is used as an additional feature for breeding Phalaenopsis. However, the scent phenotype cannot be stably inherited to the progenies, so most hybrids do not have fragrance. The assistance of molecular breeding with suitable markers for assigning to the scent loci might be practical to enhance the breeding for scent hybrids.

23.4 Phalaenopsis Breeding for Various Flower Colors

23.4.1 White Flowers

White is the most important color in Phalaenopsis. In breeding, the mean focus is on the large flowers (>10 cm) (Fig. 23.1), although the small and medium flowers and the white tepals with red lips are also becoming popular. The major features for large-and-white flowers are pure white color, large-sized, and well-rounded shape of flowers, with ordered inflorescence. Moreover, large-and-white flowers are the major breeding parents for most large-­ sized hybrids, even for the hybrids with colorful flowers. All the Phalaenopsis hybrids with large-sized flowers contain more than 50% bloodline from the large-­ and-­white flowers.

Fig. 23.1 Various phenotypes of Phalaenopsis hybrids with large-and-white flowers. (a) Phal. ‘Bataan’, (b) Phal. ‘Join Spirit’, bars=1 cm 23 Phalaenopsis 575

23.4.1.1 Native Species Contributing to Large-and-White Hybrids

The breeding of the large-and-white flowers of Phalaenopsis hybrids is the pilot in Phalaenopsis breeding history, and it has lasted for more than 100 years. The major native species contributing to the large-and-white flowers are Phal. amabilis and Phal. aphrodite, accounting for more than 90% bloodline. Phal. amabilis and Phal. aphrodite are similar to each other, and the latter has been categorized into Phal. amabilis ssp. aphrodite previously. Both of them have white flowers with the width of 7~8 cm, yellow lip with red spots, and about 30 flowers in one single inflorescence. The major differences between Phal. amabilis and Phal. aphrodite are the central lobe and callus of lip, which are deltoid and finger-like in Phal. aphrodite but oblong and shield-like in Phal. amabilis.

23.4.1.2 Important Hybrids with Large-and-White Flowers

In the breeding history of large-and-white Phalaenopsis flowers, several hybrids have played a dominant role. Phal. ‘Doris’ (Phal. ‘Elisabethae’ × Phal. ‘Katherine Siegwart’), registered in 1940, is one of these very important hybrids. Data obtained from the OrchidWiz database (2017) showed that this hybrid contributed in the development of 262 G1 (next-generation) hybrids and 30,266 hybrids when all gen- erations are taken into account (Table 23.4). A tetraploid clone of Phal. ‘Doris’ was used to breed most large-and-white hybrids with a very pure genetic background through reciprocal crosses for 15 successive generations. The most important G1 hybrids of Phal. ‘Doris’ are Phal. ‘Grace Palm’ (Phal. ‘Doris’ × Phal. ‘Winged Victory’) and its backcrossed offspring, Phal. ‘Dos Pueblos’ (Phal. ‘Doris’ × Phal. ‘Grace Palm’). These hybrids resulted in 138 and 113 of next-­generation hybrids and contributed in 24,647 and 11,826 hybrids when total generations are calculated, respectively (Table 23.4). Their progenies, Phal. ‘Juanita’ (Phal. ‘Chief Tucker’ × Phal. ‘Grace Palm’), Phal. ‘New Era’ (Phal. ‘Grace Palm’ × Phal. ‘Sally Lowrey’), Phal. ‘Ann Hatter’ (Phal. ‘Juanita’ × Phal. ‘New Era’), and Phal. ‘Pueblo Jewel’ (Phal. ‘Dos Pueblos’ × Phal. ‘Pink Jewel’), have most total numbers of hybrids derived from Phal. ‘Doris’, but only Phal. ‘Juanita’ has white flowers, while the others have white tepals with red lip, which means most of their progenies are white-tepal/red-lip hybrids (see Sect. 23.4.2.3). Other G1 hybrids of Phal. ‘Doris’, such as Phal. ‘Cast Iron Monarch’ (Phal. ‘Louise Georgianna’ × Phal. ‘Doris’), Phal. ‘Elinor Shaffer’ (Phal. ‘Juanita’ × Phal. ‘Doris’), and Phal. ‘Joseph Hampton’ (Phal. ‘Monarch Gem’ × Phal. ‘Doris’), have 69, 74, and 178 next-generation hybrids and 18,971, 8670, and 1712 hybrids of total generations, respectively, which contain white flowers with white lips (Table 23.4). Their progenies, Phal. ‘Palm Beach’ (Phal. ‘Doris’ × Phal. ‘Cast Iron Monarch’), Phal. ‘Susan Merkel’ (Phal. ‘Chieftain’ × Phal. ‘Palm Beach’), and Phal. ‘Mount Kaala’ (Phal. ‘Doreen’ × Phal. ‘Elinor Shaffer’), have also white flowers and play an important role in further development of the assortment. 576 C.-C. Hsu et al.

Table 23.4 Important hybrids for the breeding of large-and-white flowers derived fromPhal. ‘Doris’ and its progenies No. of Total no. Year next-­ of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G0 1940 (Elisabethae 262 30,266 34 9.5 WWy Doris × Katherine Siegwart) Phal. G1 1950 (Doris × 138 24,647 29 12.3 WWy Grace Winged Palm Victory) Phal. G2 1957 (Chief Tucker 87 20,470 8 12.3 WWy Juanita × Grace Palm) Phal. Dos G1 1956 (Doris × 113 11,826 5 –c WWy Pueblos Grace Palm) Phal. G2 1983 (Musashino × 50 661 3 12.2 WWy Yukimai Grace Palm) Phal. Sogo G3 1998 (Yukimai × 143 325 31 12.5 WWy Yukidian Taisuco Kochdian) Phal. Cast G1 1957 (Louise 69 18,971 3 11.5 WWy Iron Georgianna × Monarch Doris) Phal. G2 1958 (Doris × Cast 44 17,394 5 – WWy Palm Iron Beach Monarch) Phal. G3 1960 (Chieftain × 39 13,214 2 – WWy Susan Palm Beach) Merkel Phal. G1 1960 (Juanita × 74 8670 20 12.4 WWy Elinor Doris) Shaffer Phal. G2 1966 (Doreen × 98 4938 2 11.7 WWy Mount Elinor Kaala Shaffer) Phal. G1 1966 (Monarch 178 1712 9 12.1 WWy Joseph Gem × Hampton Doris) Data were collected from OrchidWiz X3.3, 2017 aFor generations. G0 is Phal. ‘Doris’. G1~G3 indicate the different generations derived from Phal. ‘Doris’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, and Wy suggests white with yellowish-like color cNot available in OrchidWiz X3.3, 2017 23 Phalaenopsis 577

Fig. 23.2 Flowers of large-and-white hybrid, Phal. ‘Sogo Yukidian’, with (a) whole plant and (b and c) flowers, bars=1 cm

In addition, the most famous hybrid within the large-and-white flowers is Phal. ‘Sogo Yukidian’ “V3” (Fig. 23.2), which is the cross of Phal. ‘Yukimai’ (seeds) to Phal. ‘Sogo Kochdian’ (pollen) by SOGO Team Co., Ltd., in Taiwan (Table 23.4). Phal. ‘Sogo Yukidian’ “V3” contains perfectly well-rounded flowers and strong and arched floral stalks with many ordered flowers. As a perfect hybrid, Phal. ‘Sogo Yukidian’ “V3” has only 143 and 308 hybrids in next and all generations, respec- tively, because the performances of its progenies are not better than itself. Therefore, “V3” almost dominates the large-and-white flowers in Phalaenopsis market. Moreover, the cross of Phal. ‘Sogo Yukidian’ “V3” and Phal. World Class “Bigfoot” provides their progeny, Phal. ‘Yu Pin Easter Island’, as the major breeding parent for Bigfoot hybrids (see Sect. 23.5.3). Genealogy analysis showed that Phal. ‘Sogo Yukidian’ “V3” contains Phal. rimestadiana (now belongs to Phal. amabilis var. rimestadiana) for 41.44%, Phal. amabilis for 40.14%, Phal. aphrodite for 15.30%, Phal. stuartiana for 2.15%, Phal. 578 C.-C. Hsu et al. schilleriana for 0.59%, and Phal. sanderiana for 0.39%. The native white species contributed for 96.88% and resulted in the large, well-rounded flowers and ordered inflorescences of Phal. ‘Sogo Yukidian’. The low percentages of Phal. stuartiana, Phal. schilleriana, and Phal. sanderiana contributed in its pure white color.

23.4.1.3 Breeding Large-and-White Flowers

In molecular genetics, the “white color” indicates “colorless,” meaning the color-­ related genes are not functional. The presence of yellow color with red spots in the lip demonstrates that the white flowers contain the color-related genes, but these are not expressed in the white tepals (Hsu et al. 2015a). Therefore, crossing white flowers with any colorful hybrid will introduce functional color-related genes and result in colorful hybrids. The major point for breeding large-and-white flowers is that only white flowers can be chosen for crossing and no colorful flowers can be used. Other important traits in breeding of the large-and-white flowers are the floral stalks with well-arched shape, ordered inflorescence, and many rounded flowers. These characteristics are already present in the bloodlines of Phal. amabilis, Phal. aphrodite, and in most of the large-and-white hybrids. However generating a better hybrid than Phal. ‘Sogo Yukidian’ “V3” is a really challenging task for breeders. The current directions for improving qualities of large-and-white flowers are mainly focused on resistance to stress caused by long distance of transport, biotic and abiotic stresses, and on plants with many floral stalks.

23.4.2 Red Flowers

The red flower hybrids comprise the highest numbers of Phalaenopsis bred off- spring. They are ranged from pink, red, lavender, to purple-red colors, as well as various pigmentation patterns, including various colors in tepals with red lip and the white flowers with red spots or stripes (called pigmentation patterning). The breeding for red flowers in Phalaenopsis can be separated into large-and-red flowers (Table 23.5, Fig. 23.3), red flowers with small and medium sizes (see Sect. 23.5.1), white tepals with red lip (Table 23.6), and white or pink tepals with red spots or stripes (Table 23.7). For the development of large-and-red flowers, Phal. ‘Doris’ and its next-generation hybrid Phal. ‘Zada’ are good breeding parents with a good performance for producing many rounded flowers and ordered inflorescences (Table 23.5). Today, two major breeding strategies within the red flower hybrids can be distinguished: large-and-red flowers and flowers containing various pigmentation patterns. 23 Phalaenopsis 579

Table 23.5 Important hybrids for the breeding of large-and-red flowers No. of Total Year next-­ no. of Natural registered generation hybrids spread Hybrid name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. Doris G0 1940 (Elisabethae × 262 30,266 34 9.5 WW Katherine Siegwart) Phal. Zada G1 1958 (San Songer × 206 15,010 9 8.4 RR Doris) Phal. G2 1968 (Ruby Wells × 139 9124 0 –c RR Lipperose Zada) Phal. G2 1969 (Doris Wells × 20 6446 0 – –d Lippezauber Zada) Phal. G3 1971 (Doris Wells × 52 5749 1 11.1 PR Lippstadt Lipperose) Phal. G3 1974 (Lippezauber 229 3723 17 10.8 PR Abendrot × Lippstadt) Phal. Flor de G2 1972 (Zada × Satin 91 2927 5 9.4 PR Mato Rouge) Phal. G3 1977 (Dear Heart × 50 2265 14 9.2 PR Herbert Flor de Mato) Hager Phal. G4 1983 (Morgenrot × 22 1734 0 – – Pinlong Herbert Cinderella Hager) Phal. New G4e 1997 (Pinlong 47 1452 2 11.7 PstR Cinderella Cinderella × New Eagle) Phal. G2 1962 (Virginia × 48 8382 2 9.8 RR Barbara Zada) Beard Phal. Lois G2f 1969 (Barbara 14 5469 2 8.9 WstR Jansen Beard × Ruby Lips) Phal. Carter G3 1972 (Lois Jansen 16 4239 0 – – Shenk × Suemid) Phal. Irene G3 1967 (Carol Brandt 10 5615 0 – – Van Alstyne × Barbara Beard) Phal. G3g 1972 (Irene Van 40 3981 1 8.5 WstR Terry-Beth Alstyne × Ballard Ruby Zada) Phal. G1 1962 (Aalsmeer 23 4456 5 8.7 PstR Raycraft Rose × Doris) Phal. G2 1973 (Grace Palm × 45 4169 0 – – Otohime Raycraft) Phal. Happy G3 1983 (Otohime × 199 3278 17 10.8 PstR Valentine Odoriko) (continued) 580 C.-C. Hsu et al.

Table 23.5 (continued) No. of Total Year next-­ no. of Natural registered generation hybrids spread Hybrid name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. Chia G4h 1982 (James Hall × 39 2919 1 6.5 WstR Lin Johanna) Phal. Ta Bei G4i 1987 (Abendrot × 39 2654 0 – – Chou Chia Lin) Phal. King G3j 1991 (Ta Bei Chou 19 1690 0 – – Shiang’s × Otohime) Coral Phal. King G5 1991 (Ta Bei Chou 3 1261 0 – – Shiang’s × Paifang’s Kide Sardonyx) Phal. King G4k 1992 (King 120 1258 2 8.7 PstR Shiang’s Shiang’s Kide Rose × King Shiang’s Coral) Data were collected from OrchidWiz X3.3, 2017 aFor generations, G0 is Phal. ‘Doris’. G1~G5 indicate the different generations derived from Phal. ‘Doris’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, Wst for white with red striped, P for pink, Pst for pink with red stripes, and R for red cNot available in OrchidWiz X3.3, 2017 dNot available in OrchidWiz X3.3, 2017 ePhal. ‘Pinlong Cinderella’ and Phal. ‘New Eagle’ are the G4 and G3 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘New Cinderella’, is the G4 progenies of Phal. ‘Doris’ fPhal. ‘Barbara Beard’ and Phal. ‘Ruby Lips’ are the G2 and G1 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘Lois Jansen’, is the G2 progenies of Phal. ‘Doris’ gPhal. ‘Irene Van Alstyne’ and Phal. ‘Ruby Zada’ are the G3 and G2 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘Terry-Beth Ballard’, is the G3 progenies of Phal. ‘Doris’ hPhal. ‘Chia Lin’ is the G4 progenies of Phal. ‘Doris’ from the cross of G5 Phal. ‘James Hall’ (Phal. ‘Red Lip’ × Phal. ‘Barbara Moler’) to G3 Phal. ‘Johanna’ (Phal. ‘Ella Freed’ × Phal. ‘Jiminy Cricket’) iPhal. ‘Abendrot’ and Phal. ‘Chia Lin’ are the G3 and G4 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘Ta Bei Chou’, is the G4 progenies of Phal. ‘Doris’ jPhal. ‘Ta Bei Chou’ and Phal. ‘Otohime’ are the G4 and G2 progenies of Phal. ‘Dori's, respec- tively, so their hybrid, Phal. ‘Ta Bei Chou’, is the G3 progenies of Phal. ‘Doris’ kPhal. ‘King Shiang’s Coral’ and Phal. ‘King Shiang’s Kide’ are the G3 and G5 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘King Shiang’s Rose’, is the G4 progenies of Phal. ‘Doris’

23.4.2.1 Native Species Contributing to Red Flower Hybrids

For large-and-red flowers, Phal. schilleriana and Phal. sanderiana are the major native species used as breeding parents. Phal. schilleriana contains scented flowers, 5~8 cm in width, pink to red colors, many-flower inflorescences, and branched floral stalks (Fig. 23.4). Phal. sanderiana has 5~8 cm flowers with pink to 23 Phalaenopsis 581

Fig. 23.3 Various phenotypes of Phalaenopsis hybrids with large-and-red flowers. (a) Phal. ‘Taisuco Peace’, (b) Phal. ‘Nobby’s Spring Alice’, (c) Phal. ‘Taida New Luchia’, (d) Phal. ‘Hong Lin Jewelry’, (e) Phal. ‘OX Pink Yukidian’, (f) Phal. ‘Dragon Tree Firerose’, bars=1 cm

purple-­red colors and many-flower inflorescences. Due to the introduction of Phal. ‘Doris’, the bloodlines of Phal. amabilis and Phal. aphrodite are also important for large-­and-­red flowers. In addition, Phal. equestris and Phal. pulcherrima are the major wild species for breeding of red hybrids with small and medium sizes. Phal. equestris contains 2~4 cm flowers with various colors, including white, pink, red, purple-to-red, and purple-to-blue colors. Phal. equestris has branched and many-flower inflorescences 582 C.-C. Hsu et al.

Table 23.6 Important hybrids for the breeding of white tepals and red lip No. of Total no. Year next-­ of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G0 1940 (Elisabethae 262 30,266 34 9.5 WW Doris × Katherine Siegwart) Phal. –c 1954 (Pua Kea × 32 17,085 2 –d –e Sally equestris) Lowrey Phal. G2 1957 (Chieftain × 17 12,785 3 9.5 WR Judy Sally Karleen Lowrey) Phal. G2 1958 (Grace Palm 9 14,378 2 – WR New Era × Sally Lowrey) Phal. Ann G3 1962 (Juanita × 21 13,531 4 9.8 WR Hatter New Era) Phal. G1 1958 (Sally 4 136 1 8.0 WO Cover Lowrey × Girl Doris) Phal. G2 1968 (Dos Pueblos 22 6588 1 8.9 WR Pueblo × Pink Jewel) Jewel Phal. G4 1991 (Pamela Lady 7 222 1 11.0 WR Luchia × Pinlong Lady Cardinal) Phal. G4 1993 (Musashino × 8 135 1 11.9 WR Hsinying Su’s Red Lip Lip) Phal. G5 1998 (Luchia 32 106 3 10.0 WR Luchia Lady × Lip Hsinying Lip) Phal. G3 1993 (Mount 26 766 0 – – Mount Kaala × Beauty Hamakita Beauty) Phal. Su’s – 1986 (South 70 950 1 9.5 WR Red Lip Cha-Li × Lucky Lady) Phal. G4 1993 (Mount 7 10 0 – WR Hsin Red Beauty × Lip Su’s Red Lip) (continued) 23 Phalaenopsis 583

Table 23.6 (continued) No. of Total no. Year next-­ of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G3 1997 (South 88 348 8 10.4 WR Mount Cha-Li × Lip Mount Beauty) Phal. G4 2000 (Mount Lip 18 64 4 10.6 WR Hsinying × Tinny Ace) Mount Data were collected from OrchidWiz X3.3, 2017 aFor generations, G0 is Phal. ‘Doris’. G1~G5 indicate the different generations derived from Phal. ‘Doris’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, O for orange, P for pink, R for red cNot the progenies of G0, Phal. ‘Doris’ dNot available in OrchidWiz X3.3, 2017 eNot available in OrchidWiz X3.3, 2017 and many floral stalks.Phal. pulcherrima has 2~3 cm flowers with various colors, including white, pink, purple-to-red, deep-red, and purple-to-blue colors. Moreover, other native species provide different color traits, like the various pig- mentation patterning. Phal. stuartiana has white flowers with red spots and is the major breeding parent for red-spot feature. Phal. lindenii introduces the red-stripe trait, and Phal. pulcherrima gives the dark-red and purple colors.

23.4.2.2 Important Hybrids with Large-and-Red Flowers

The cross between the white-flowerPhal. ‘Doris’ (pollen) and the red-flower Phal. ‘San Songer’ (seeds) resulted in 1958 in the introduction of Phal. ‘Zada’, which became the major breeding parent for large red flowers with 206 and 15,010 hybrids in next-generation and all generations, respectively (Table 23.5). Its next-generation hybrids, Phal. ‘Lipperose’ (Phal. ‘Ruby Wells’ × Phal. ‘Zada’), Phal. ‘Flor de Mato’ (Phal. ‘Zada’ × Phal. ‘Satin Rouge’), and Phal. ‘Barbara Beard’ (Phal. ‘Virginia’ × Phal. ‘Zada’) are the most used breeding parents (Table 23.5). Phal. ‘Lipperose’ belongs to a series of Phalaenopsis hybrids with large red flowers and were registered by Hark-Orchideen in Germany in the 1970s. More hybrids in this series include Phal. ‘Lippezauber’ (Phal. ‘Doris’ Wells × Phal. ‘Zada’), Phal. ‘Lippstadt’ (Phal. ‘Doris Wells’ × Phal. ‘Lipperose’), and Phal. ‘Abendrot’ (Phal. ‘Lippezauber’ × Phal. ‘Lippstadt’) which were used frequently as breeding parents. All these cultivars had flowers with 10.8~11.1 cm width and pink to red tepals with a red lip. Among them, Phal. ‘Abendrot’ was most used as hybrid- ization parent (Table 23.5). 584 C.-C. Hsu et al.

Table 23.7 Important hybrids for the breeding of flowers with red stripes (venation) or red spots No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. –c 1960 (lindenii × 2 3 0 –d –e Robert sanderiana) W. Miller Phal. – 1964 (lindenii × Pink 40 81 6 4.7 PstR Peppermint Profusion) Phal. – 1966 (schilleriana × 15 118 2 – WstP Baguio lindenii) Phal. Doris G0 1940 (Elisabethae × 262 30,266 34 9.5 WW Katherine Siegwart) Phal. Star – 1956 (Bataan × 13 10,635 4 10.2 WspR of Rio lueddemanniana) Phal. – 1963 (Star of Rio × 87 7735 20 7.3 WspR Samba amboinensis) Phal. Ella G2 1970 (Show Girl × 129 7268 8 9.8 WstR Freed Samba) Phal. G3 1975 (Ella Freed × 54 4180 5 9.0 WstR Freed’s Career Girl) Danseuse Phal. G4 1987 (Freed’s 21 3513 0 – – Modern Danseuse × Stripes Chiali Stripe) Phal. Okay G5 1992 (Modern Stripes 24 2170 1 8.2 WstR Seven × Houpi Beauty) Phal. G6 1994 (Okay Seven × 13 318 2 6.7 WstR Taisuco Taisuco Gaster) Stripe Phal. Little G7 1997 (Taisuco Stripe 144 292 16 5.4 WstR Gem × Taisuco Gem) Stripes Phal. G4 1979 (Lucky Lady × 4 4183 0 – – Lucky Carter Shenk) Shenk Phal. Chiali G4f 1983 (Cindy Tsai × 21 4175 0 – – Stripe Lucky Shenk) Phal. G5 1987 (Tsuei You 5 3441 0 – – Houpi Queen × Chiali Beauty Stripe) Phal. Sun G5g 1995 (Cypress Pink × 39 2148 0 – – Prince Houpi Beauty) Phal. G6 1997 (Sun Prince × 221 567 32 9.3 WspR Leopard Ho’s French Prince Fantasia) (continued) 23 Phalaenopsis 585

Table 23.7 (continued) Data were collected from OrchidWiz X3.3, 2017 aFor generations, G0 is Phal. ‘Doris’. G1~G7 indicate the different generations derived from Phal. ‘Doris’ bThe colors for cultivar flowers. First one is the color for tepals, and second one is the color for lip. Wst indicates white with red stripes, Wsp for white with red spots, O for orange, P for pink, R for red cNot the progenies of G0, Phal. ‘Doris’ dNot available in OrchidWiz X3.3, 2017 eNot available in OrchidWiz X3.3, 2017 fPhal. ‘Cindy Tsai’ and Phal. ‘Lucky Shenk’ are the G3 and G4 progenies of Phal. ‘Doris’, respec- tively, so their hybrid, Phal. ‘Chiali Stripe’, is the G4 progenies of Phal. ‘Doris’ gPhal. ‘Cypress Pink’ and Phal. ‘Houpi Beauty’ are the G4 and G5 progenies of Phal. ‘Doris’, respectively, so their hybrid, Phal. ‘Sun Prince’, is the G5 progenies of Phal. ‘Doris’

Fig. 23.4 Flowers of red Phalaenopsis species, Phal. schilleriana, bars=1 cm

The progenies of Phal. ‘Flor de Mato’, Phal. ‘Herbert Hager’ (Phal. ‘Dear Heart’ × Phal. ‘Flor de Mato’), Phal. ‘Pinlong Cinderella’ (Phal. ‘Morgenrot’ × Phal. ‘Herbert Hager’), and Phal. ‘New Cinderella’ (Phal. ‘Pinlong Cinderella’ × Phal. ‘New Eagle’) were the serial hybrids containing most total progenies with 50, 22, and 47 next-generation hybrids and 2265, 1734, and 1452 progenies of total genera- tions, respectively. Phal. ‘Herbert Hager’ had 9.2 cm large flowers with pink tepals and red lip, while Phal. ‘New Cinderella’ had 11.7 cm flowers and pink tepals with red stripes and red lip. Another important breeding line originated from Phal. ‘Barbara Beard’, Phal. ‘Lois Jansen’ (Phal. ‘Barbara Beard’ × Phal. ‘Ruby Lips’), and Phal. ‘Irene Van Alstyne’ (Phal. ‘Carol Brandt’ × Phal. ‘Barbara Beard’) and their next-generation hybrids, Phal. ‘Carter Shenk’ (Phal. ‘Lois Jansen’ × Phal. ‘Suemid’) and Phal. ‘Terry-Beth­ Ballard’ (Phal. ‘Irene Van Alstyne’ × Phal. ‘Ruby Zada’) (Table 23.3). All these hybrids have similar flower sizes of 8.5~9.8 cm, but Phal. ‘Barbara Beard’ has red flowers, while Phal. ‘Lois Jansen’ and Phal. ‘Terry-Beth Ballard’ have white to pink tepals with red strips. Except from the bloodline of Phal. ‘Zada’, Phal. ‘Raycraft’ (Phal. ‘Aalsmeer Rose’ × Phal. ‘Doris’) and its progenies, Phal. ‘Otohime’ (Phal. ‘Grace Palm’ × Phal. 586 C.-C. Hsu et al.

‘Raycraft’) and Phal. ‘Happy Valentine’ (Phal. ‘Otohime’ × Phal. ‘Odoriko’), had contributed significantly in the development of the red flower assortment. These hybrids have the flowers with 8,7~10.8 cm in size and pink tepals containing red stripes and red lip. The introduction of the bloodline of Phal. pulcherrima into red-flower breeding was started from Phal. ‘Red Coral’ (Phal. pulcherrima var. buyssoniana × Phal. ‘Doris’) and Phal. ‘Memoria Clarence Schubert’ (Phal. pulcherrima var. buyssoni- ana × Phal. ‘Zada’), which are used as breeding parents for 81 and 62 next-genera- tion hybrids and 289 and 221 hybrids of total generations, respectively. These two hybrids have similar flower sizes about 6.5~7.6 cm and a red color. Phal. ‘Chia Lin’ (Phal. ‘James Hall’ × Phal. ‘Johanna’) is the fourth generation from Phal. ‘Doris’ and contains two different colors, purple-red flowers or yellow tepals with red lip (Table 23.5). So Phal. ‘Chia Lin’ is a major parent for breeding both the purple-red flowers and the yellows tepals with red lip (see Sect. 23.4.3.3). Phal. ‘Chia Lin’ had 39 next-generation hybrids and 2919 hybrids in all genera- tions, but most of the next-generation hybrids contained yellow and middle-sized flowers, except for Phal. ‘Ta Bei Chou’ (Phal. ‘Abendrot’ × Phal. ‘Chia Lin’), Phal. ‘Brother Cardinal’ (Phal. ‘Chia Lin’ × Phal. ‘Pinlong Major’), and Phal. ‘Taida David’ (Phal. ‘Chia Lin’ × Phal. ‘Sogo David’). Among them, Phal. ‘Ta Bei Chou’ has most progenies, and its offspring, Phal. ‘King Shiang’s Coral’ (Phal. ‘Ta Bei Chou’ × Phal. ‘Otohime’), Phal. ‘King Shiang’s Kide’ (Phal. ‘Ta Bei Chou’ × Phal. ‘Paifang’s Sardonyx’), and Phal. ‘King Shiang’s Rose’ (Phal. ‘King Shiang’s Kide’ × Phal. ‘King Shiang’s Coral’), are major breeding parents for red flowers. The flowers of Phal. ‘King Shiang’s Rose’ are 8.7 cm in width and are characterized with pink tepals with red stripes and red lip.

23.4.2.3 Important Hybrids with White Tepals and Red Lip

Most Phalaenopsis hybrids with flowers existing of white tepals with a red lip are derived from small-and-red Phal. equestris and yellow-flower/red-lipPhal. luedd- emanniana. For example, Phal. ‘Sally Lowrey’ (Phal. ‘Pua Kea’ × Phal. equestris) has small-and-pink flowers and was used to cross with large-and-white flowers to increase the flower size (Table 23.6). Therefore, Phal. ‘Judy Karleen’ (Phal. ‘Chieftain’ × Phal. ‘Sally Lowrey’), Phal. ‘New Era’ (Phal. ‘Grace Palm’ × Phal. ‘Sally Lowrey’), and Phal. ‘Cover Girl’ (Phal. ‘Sally Lowrey’ × Phal. ‘Doris’) have white tepals and red lip and were used further as breeding parents for 17, 9, and 4 next-generation hybrids and 12,785, 14,378, and 136 hybrids of total generations, respectively. In Phalaenopsis market, the most famous flowers with white tepals and red lip are Phal. ‘Hsin Red Lip’ (Phal. ‘Mount Beauty’ × Phal. ‘Su’s Red Lip’), Phal. ‘Luchia Lip’ (Phal. ‘Luchia Lady’ × Phal. ‘Hsinying Lip’), Phal. ‘Mount Lip’ (Phal. ‘South Cha-Li’ × Phal. ‘Mount Beauty’), and Phal. ‘Hsinying Mount’ (Phal. ‘Mount Lip’ × Phal. ‘Tinny Ace’) (Table 23.6). Phal. ‘Luchia Lip’ was hybridized from the cross of Phal. lueddemanniana to large-and-white flowers for several generations to produce large flowers with white tepals and red lip, and the chosen breeding line of Phal. 23 Phalaenopsis 587

Fig. 23.5 Flowers of white tepals and red lip. (a) Phal. ‘Mount Lip’, (b) Phal. ‘Fuller’s Pink Rose’, bars=1 cm

‘Luchia Lip’ has 10 cm flowers and resulted in 32 next-generation and 106 hybrids of total generations. So does Phal. ‘Hsin Red Lip’, which was bred from the cross of both white-tepal/red-lip flowers of Phal. ‘Su’s Red Lip’ (pollen) and Phal. ‘Mount Beauty’ (seeds). Phal. ‘Mount Lip’ (Fig. 23.5) and Phal. ‘Hsinying Mount’ are famous for their flowers having white tepals with red lip and the pink color at the base of tepals that make the white flowers contain red color in the flower center. Phal. ‘Mount Lip’ and Phal. ‘Hsinying Mount’ were further used in breeding (Table 23.6). Genealogy analysis showed that Phal. ‘Mount Lip’ contains Phal. rimestadiana for 37.0%, Phal. amabilis for 35.4%, Phal. aphrodite for 17.2%, Phal. equestris for 3.5%, Phal. schilleriana for 2.0%, Phal. sanderiana for 1.9%, Phal. lueddemanniana for 1.6%, and others.

23.4.2.4 Important Hybrids with Red-Stripe Flowers

The feature of flowers with red stripes came from a native species, Phal. lindenii, which has white flowers with red stripes accompanied with veins, so-called vena- tion pattern. Phal. lindenii has been used as breeding parent for red-stripe trait, such as Phal. ‘Robert W. Miller’ (Phal. lindenii × Phal. sanderiana), Phal. ‘Peppermint’ (Phal. lindenii × Phal. ‘Pink Profusion’), and Phal. ‘Baguio’ (Phal. schilleriana × Phal. lindenii) (Table 23.7). Most hybrids with red stripes came from the red-spot flowers, where the red spots lined along with veins and looks like the stripes. For example, Phal. ‘Star of Rio’ (Phal. ‘Bataan’ × Phal. lueddemanniana) came from the cross of white-flower/ red-stripe Phal. lueddemanniana and white-flowerPhal. ‘Bataan’ (Phal. amabilis × Phal. aphrodite) and has the red spots radiating from the center of tepals. Its next-­ generation hybrid, Phal. ‘Samba’ (Phal. ‘Star of Rio’ × Phal. amboinensis), has also red spots in white flowers. Phal. ‘Star of Rio’ and Phal. ‘Samba’ were further used to generate new hybrids. However, their progenies, Phal. ‘Ella Freed’ (Phal. ‘Show Girl’ × Phal. ‘Samba’), Phal. ‘Freed’s Danseuse’ (Phal. ‘Ella Freed’ × Phal. 588 C.-C. Hsu et al.

Fig. 23.6 Various phenotypes of Phalaenopsis hybrids with white color and red stripes. (a) Phal. ‘Little Gem Stripes’, (b) Phal. ‘Taida Salu’, (c) Phal. ‘Taisuco Glitter’, (d) Phal. ‘Fuller’s Pink Gem’, (e) Phal. ‘OX Leo Prince’, (f) Phal. ‘I-Hsin The Big Bang’, bars=1 cm

‘Career Girl’), Phal. ‘Modern Stripes’ (Phal. ‘Freed’s Danseuse’ × Phal. ‘Chiali Stripe’), Phal. ‘Okay Seven’ (Phal. ‘Modern Stripes’ × Phal. ‘Houpi Beauty’), Phal. ‘Taisuco Stripe’ (Phal. ‘Okay Seven’ × Phal. ‘Taisuco Gaster’), and Phal. ‘Little Gem Stripes’ (Phal. ‘Taisuco Stripe’ × Phal. ‘Taisuco Gem’), all have pink to red stripes in white tepals and red lip (Table 23.7). Among them, Phal. ‘Little Gem Stripes’ (Fig. 23.6) is still usually used as breeding parent for both white and yellow flowers with red stripes. For introducing the large-flower trait, the large red hybrid, Phal. ‘Carter Shenk’, was used to cross with Phal. ‘Lucky Lady’ and resulted in Phal. ‘Lucky Shenk’ 23 Phalaenopsis 589

(Phal. ‘Lucky Lady’ × Phal. ‘Carter Shenk’). Subsequently the hybrid was further used for the generation of Phal. ‘Chiali Stripe’ (Phal. ‘Cindy Tsai’ × Phal. ‘Lucky Shenk’), Phal. ‘Houpi Beauty’ (Phal. ‘Tsuei You Queen’ × Phal. ‘Chiali Stripe’), Phal. ‘Sun Prince’ (Phal. ‘Cypress Pink’ × Phal. ‘Houpi Beauty’), and Phal. ‘Leopard Prince’ (Phal. ‘Sun Prince’ × Phal. ‘Ho’s French Fantasia’). In addition, Phal. ‘Leopard Prince’ has white flowers with red stripes and won 32 awards. Phal. ‘Leopard Prince’ has been used as breeding parent for 221 next-generation hybrids and 567 hybrids of total generations (Table 23.7). Genealogy analysis showed that Phal. ‘Little Gem Stripes’ contains Phal. eques- tris for 29.3%, Phal. amabilis for 21.7%, Phal. rimestadiana for 15.9%, Phal. aph- rodite for 9.6%, Phal. schilleriana for 7.9%, Phal. sanderiana for 5.3%, Phal. lueddemanniana for 4.4%, Phal. amboinensis for 4.3%, Phal. stuartiana for 1.0%, and others.

23.4.2.5 Molecular Genetics for Breeding Red Flowers

Anthocyanins are water-soluble pigments that occur in almost all plants and are responsible for most of orange, red, purple, and blue colors of flowers. The biosynthetic pathway for anthocyanin accumulation is one of the most extensively studied in plant secondary metabolism, including the genes of chalcone synthase (CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3′-hydroxylase (F3′H), flavonoid 3′5′-hydroxylase (F3′5′H), dihydroflavonol 4-reductase (DFR), anthocyanidin synthase (ANS), and UDP-glucose:flavonoid 3-O-glucosyltransferase (UFGT) (Grotewold 2006). In addition, many regulatory genes involved in the anthocyanin biosynthesis have also been cloned and characterized from a wide variety of plants. R2R3-MYB and basic helix-loop-helix (bHLH) transcription factors and WD40 repeat (WDR) proteins are three major families of regulatory proteins for anthocyanin biosynthesis (Winkel-Shirley 2001; Koes et al. 2005; Feller et al. 2011; Hichri et al. 2011; Petroni and Tonelli 2011). Among them, these R2R3-MYB transcription factors are key components providing the specificity for the downstream genes and causing tissue-specific anthocyanin accumulation (Borevitz et al. 2000; Zhang et al. 2003). In Phal. schilleriana, PsDFR and PsMYB were expressed in its purple flowers and the black spots of petals of Phal. ‘Ever-spring Fairy’ (Ma et al. 2009). PeUFGT3 (Phal. equestris UFGT3) is highly expressed in the red lip, but not in the white tepals, of Phal. ‘Luchia Lady’, and other red-flower hybrids (Chen et al. 2011). Whole-genome screening for the genes participated in anthocyanin biosynthesis pathway has been performed and found that three structural genes, PeF3H5, PeDFR1, and PeANS3, and three regulatory genes, PeMYB2, PeMYB11, and PeMYB12, regulate anthocyanin accumulation in Phalaenopsis (Hsu et al. 2015a). Moreover, these three PeMYBs regulate distinct pigmentation patterns in one single flower, and the regulations of pigmentation patterning by PeMYBs are very different in tepals and lip. In tepals, PeMYB2, PeMYB11, and PeMYB12 control the full-red pigmentation, red spots, and venation pattern, respectively. In lip, PeMYB11 590 C.-C. Hsu et al. regulates the red spots in the callus, and PeMYB12 is the major regulator for pigmentation in the central lobe. Combination of the regulatory effects of these three PeMYBs leads to a very complicated floral pigmentation patterning in Phalaenopsis (Hsu et al. 2015a).

23.4.3 Yellow Flowers

The Phalaenopsis hybrids with yellow color only occupy a small portion of Phalaenopsis market. However, the bright feeling of yellow color and the rare number of hybrids make the breeding for yellow-flower hybrids also important. The yellow-colored hybrids can be subdivided in flowers with pure yellow (Table 23.8, Fig. 23.7), yellow tepals with red lip, and yellow flowers with red stripes (Table 23.8) as well as orange and green colors (Table 23.9). The major problem for breeding yellow flowers is the color decay after the flower opening, even to nearly white color. So for a good hybrid, the yellow color needs to be bright or dense and lasts for all flowering stages. In addition, most yellow-flower hybrids contain one floral stalk with few flowers, and the flowers have the star-like shape due to the wild species with yellow flowers. So the yellow-flower hybrids need to be crossed with white-flower hybrids to change the features of the flower morphology and to obtain well-rounded flowers.

23.4.3.1 Native Species Contributing to Yellow-Flower Hybrids

Among the 92 native species, only about 20 species contain yellow flower color, and these all belong to subgenus Polychilos. Among them, Phal. amboinensis, Phal. cornu-cervi, Phal. fasciata, Phal. gigantea, Phal. lueddemanniana, Phal. mannii, Phal. sumatrana, and Phal. venosa are most used (Fig. 23.8). These native species with yellow flowers have mostly few and star-like flowers and therefore need to be crossed with white flowers to increase the flower numbers and to create better flower morphology. However, due to differences in chromosome size between the subgenus Polychilos, to which most yellow-flowered species belong, and the large white flow- ering species of subgenus Phalaenopsis, crosses between these two subgenera are very challenging to succeed, and mostly sterile hybrids are obtained (see Sect. 23.7).

23.4.3.2 Important Hybrids with Yellow Flowers

The first famous yellow-flower hybrid is Phal. ‘Golden Emperor’ (Phal. ‘Snow Daffodil’ × Phal. ‘Mambo’), which contains well-rounded flower shape and pure yellow color. It was registered in 1982. However, the chromosome number of Phal. ‘Golden Emperor’ is 2n = 3x, so progenies of Phal. ‘Golden Emperor’ are very rare and only 7 hybrids derived. 23 Phalaenopsis 591

Table 23.8 Important hybrids for the breeding of yellow flowers from Phal. ‘Golden Sands’, Phal. ‘Taipei Gold’, and their progenies No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Yellow flowers Phal. G0 1964 (Fenton Davis 137 6572 18 8.4 YspO Golden Avant × Sands lueddemanniana) Phal. G1 1976 (Golden Sands × 113 2845 11 7.0 YspO Golden amboinensis) Amboin Phal. Salu G2 1992 (Paifang’s 44 1467 3 7.7 YspO Spot Auckland × Golden Amboin) Phal. G3 1995 (Salu Spot × 102 1099 5 8.0 YspO Golden Sentra) Sun Phal. G1 1979 (Golden Sands × 19 71 29 6.8 YspO Solar Golden Pride) Flare Phal. Liu G1 1979 (gigantea × 80 4528 6 8.0 YspO Tuen-­ Golden Sands) Shen Phal. G2 1983 (Liu Tuen-Shen 16 44 3 7.6 GR Fortune × Barbara Freed Saltzman Saltzman) Phal. G2 1983 (Misty Green × 208 3227 9 7.1 WspO Golden Liu Tuen-Shen) Peoker Phal. G0 1984 (Gladys Read × 152 1776 14 8.0 YspR Taipei venosa) Gold Phal. G1 1995 (Taipei Gold × 108 762 27 7.7 YspR Brother Deventeriana) Lawrence Phal. G1 1995 (Taipei Gold × 8 192 0 –c YspR Brother Brother Imp) Nugget Phal. G1 2001 (Taipei Gold × 77 182 2 7.0 YspR I-Hsin Brother Nugget) Sunflower Phal. G2 1997 (Brother 92 455 12 6.8 YspR Sogo Lawrence × Manager Autumn Sun) (continued) 592 C.-C. Hsu et al.

Table 23.8 (continued) No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G2 2000 (Brother 87 172 5 6.4 YspR Sogo Lawrence × Pride Sogo Manager) Yellow tepals and red lip Phal. G4 1982 (James Hall × 39 2919 1 6.5 WstR Chia Lin Johanna) Phal. G5 2003 (Chia Lin × 58 102 0 – YR Sunrise Tinny Honey) Star Phal. G5 2004 (Chia Lin × 44 170 0 – YstR Chian Mount Beauty) Xen Queen Phal. G6 2004 (Taisuco Date × 24 26 17 8.9 YR Fuller’s Chian Xen Sunset Queen) Yellow flowers with red stripes Phal. –d 1997 (Salu Spot × 29 38 8 7.8 YstR Taida Happy Beauty) Salu Phal. – 1997 (Taisuco Stripe × 144 292 16 5.4 WstR Little Taisuco Gem) Gem Stripes Phal. – 2004 (Little Gem 0 0 0 – –e Taida Stripe × Salu Golden Peoker) Gem Phal. – 2009 (Little Gem 5 5 15 6.1 YstR Fangmei Stripe × Dou-dii Sweet Golden Princess) Data were collected from OrchidWiz X3.3, 2017 aFor generations, G0 is Phal. ‘Golden Sands’ or Phal. ‘Taipei Gold’. G1~G6 indicate the different generations derived from Phal. ‘Golden Sands’ or Phal. ‘Taipei Gold’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. Wst indicates white with stripes, Wst for white with spots, G for yellowish green, O for orange, R for red, Y for yellow, Ysp for yellow with red spots, and Yst for yellow with red stripes cNot available in OrchidWiz X3.3, 2017 dNot the progenies of G0, Phal. ‘Golden Sands’ or Phal. ‘Taipei Gold’ eNot available in OrchidWiz X3.3, 2017 23 Phalaenopsis 593

Fig. 23.7 Various phenotypes of Phalaenopsis hybrids with yellow flowers. (a) Phal. ‘Chian Xen Gold’, (b) Phal. ‘Golden Apollon’, (c) Phal. ‘I-Hsin Bright Star’, (d) Phal. ‘Taisuco Date’, bars=1 cm

Phal. ‘Golden Sands’ (Phal. ‘Fenton Davis Avant’ × Phal. lueddemanniana) brought the bloodline of Phal. lueddemanniana into yellow-flower breeding and is a major breeding parent for 137 next-generation hybrids and 6572 hybrids of total generations (Table 23.8). Phal. ‘Golden Sands’ and most of its progenies contain yellow flowers with red spots and orange lip. Its progenies, Phal. ‘Golden Amboin’ (Phal. ‘Golden Sands’ × Phal. amboinensis), Phal. ‘Salu Spot’ (Phal. ‘Paifang’s Auckland’ × Phal. ‘Golden Amboin’), and Phal. ‘Golden Sun’ (Phal. ‘Salu Spot’ × Phal. ‘Sentra’), are important breeding parents. Among the total progenies of Phal. ‘Golden Sands’, the G1-hybrid Phal. ‘Solar Flare’ (Phal. ‘Golden Sands’ × Phal. ‘Golden Pride’) and G4-hybrid Phal. ‘Ambo Buddha’ (Phal. ‘Brother Buddha’ × Phal. amboinensis) won the most awards. Genealogy analysis showed that Phal. ‘Ambo Buddha’ contains Phal. amboi- nensis for 56.3%, Phal. rimestadiana for 7.3%, Phal. fasciata for 6.3%, Phal. gigantea for 6.3%, Phal. lueddemanniana for 6.3%, Phal. amabilis for 4.0%, Phal. aphrodite for 3.2%, Phal. equestris for 2.3%, and Phal. sanderiana for 1.2%. The cross between Phal. ‘Golden Sands’ (pollen) and the wax-textured flowers of Phal. gigantea (seeds) produced Phal. ‘Liu Tuen-Shen’, which has flowers with better texture and was further used as breeding parent for 80 G1 hybrids and 4528 hybrids in all generations (Table 23.8). From its progeny, Phal. ‘Golden Peoker’ 594 C.-C. Hsu et al.

Table 23.9 Important hybrids for the breeding of orange and green flowers No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Orange flowers Phal. – 1961 (violacea × 225 2545 29 –c RR Princess amboinensis) Kaiulani Phal. – 1895 (lueddemanniana 82 2197 31 5.3 RR Luedde-­ × violacea) violacea Phal. – 1974 (violacea × 128 1692 11 5.0 RR George Luedde-violacea) Vasquez Phal. – 1997 (Sara Lee × 91 351 13 6.0 OspR Brother Taipei Gold) Sara Gold Phal. Salu – 2000 (Golden Sun × 86 289 4 7.2 OspR Peoker Yellow Peoker) Phal. KV – 2006 (Salu Peoker × 6 7 4 6.3 OspR Golden Brother Sara Star Gold) Phal. Sogo – 2002 (Brother Peter 66 161 7 5.6 OspR Lawrence Star × Brother Sara Gold) Phal. Surf – 2004 (Taipei Gold × 14 14 3 7.9 OR Song King Shiang’s Rose) Green flowers Phal. – 1984 (amboinensis × 36 1064 1 6.1 GspW Gelblieber micholitzii) Phal. – 1995 (Gelblieber × 86 296 4 5.1 YY Yungho Princess Gelb Kaiulani) Canary Phal. – 1986 (Gelblieber × 45 270 4 4.2 GW Su-An mannii) Cricket Phal. – 1994 (Gelblieber × 41 140 2 5.8 GspW Hannover mariae) Passion Phal. – 2005 (Gelblieber × 10 13 17 5.5 GW Nobby’s Nobby’s Fox) Green Eagle (continued) 23 Phalaenopsis 595

Table 23.9 (continued) No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. – 1983 (Liu Tuen-Shen 16 44 3 7.6 GR Fortune × Barbara Freed Saltzman Saltzman) Data were collected from OrchidWiz X3.3, 2017 aNo common parents bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, G for yellowish green, Gsp for greenish with spots, O for orange, Osp for orange with spots, P for pink, R for red, and Y for yellow cNot available in OrchidWiz X3.3, 2017

Fig. 23.8 These native species for breeding yellow Phalaenopsis hybrids. (a) Phal. lueddeman- niana, (b) Phal. amboinensis, (c) Phal. gigantea, (d) Phal. mannii, bars=1 cm 596 C.-C. Hsu et al.

Fig. 23.9 Various phenotypes of Phalaenopsis hybrids with yellow tepals and red lip. (a) Phal. ‘Taisuco Gloria’, (b) Phal. ‘Fusheng’s Golden Age’, bars=1 cm

(Phal. ‘Misty Green’ × Phal. ‘Liu Tuen-Shen’) started the breeding of harlequin flowers by the selection of an “accident” mutant from somaclonal variation (see Sect. 23.4.4). Another progeny, Phal. ‘Fortune Saltzman’ (Phal. ‘Liu Tuen-Shen’ × Phal. ‘Barbara Freed Saltzman’), has green tepals with red lip (see Sect. 23.4.3.6). The introduction of Phal. venosa into the yellow-flower breeding started from Phal. ‘Taipei Gold’ (Phal. ‘Gladys Read’ × Phal. venosa) in 1984, which have 152 next-generation hybrids and 1776 hybrids of total generations. Among them, Phal. ‘Brother Lawrence’ (Phal. ‘Taipei Gold’ × Phal. ‘Deventeriana’), Phal. ‘Brother Nugget’ (Phal. ‘Taipei Gold’ × Phal. ‘Brother Imp’), Phal. ‘I-Hsin Sunflower’ (Phal. ‘Taipei Gold’ × Phal. ‘Brother Nugget’), Phal. ‘Sogo Manager’ (Phal. ‘Brother Lawrence’ × Phal. ‘Autumn Sun’), and Phal. ‘Sogo Pride’ (Phal. ‘Brother Lawrence’ × Phal. ‘Sogo Manager’) were major breeding parents.

23.4.3.3 Important Hybrids with Yellow Flowers with Red Lip

Phal. ‘Chia Lin’ (Phal. ‘James Hall’ × Phal. ‘Johanna’) is a major parent for breed- ing both the purple-red flowers and yellow tepals with red lip (Table 23.8). Most of its G1 hybrids have yellow tepals with red lip, and few of them contain red-to-pur- ple flowers. Among them, Phal. ‘Sunrise Star’ (Phal. ‘Chia Lin’ × Phal. ‘Tinny Honey’) and Phal. ‘Chian Xen Queen’ (Phal. ‘Chia Lin’ × Phal. ‘Mount Beauty’) are famous yellow-flower hybrids with red lip. Phal. ‘Chian Xen Queen’ has a very famous G1 hybrid, Phal. ‘Fuller’s Sunset’ (Phal. ‘Taisuco Date’ × Phal. ‘Chian Xen Queen’), which has bright yellow tepals with red lip, and had 24 and 26 hybrids of next and total generations, respectively. Phal. ‘Fusheng’s Golden Age’ (Phal. ‘Fong-Tien’s Yellow Butterfly’ × Phal. ‘Chian Xen Queen’) is also a next-genera- tion hybrid of Phal. ‘Chian Xen Queen’ and has bright yellow tepals and red lip (Fig. 23.9). Genealogy analysis showed that Phal. ‘Fuller’s Sunset’ contains Phal. amabilis for 28.0%, Phal. rimestadiana for 19.8%, Phal. amboinensis for 14.1%, 23 Phalaenopsis 597

Fig. 23.10 Various phenotypes of Phalaenopsis hybrids with yellow color with red stripes. (a) Phal. ‘Fangmei Sweet’, (b) Phal. ‘Fuller’s Gold Stripes’, bars=1 cm

Phal. aphrodite for 9.7%, Phal. venosa for 9.4%, Phal. lueddemanniana for 5.5%, and others. The yellow color might be from the bloodline of Phal. venosa, and the red color in lip might be resulted from Phal. amboinensis.

23.4.3.4 Important Hybrids for Yellow Flowers with Red Stripes

Most yellow flowers with red stripes came from the cross between the yellow flow- ers with red spots and the white flowers with red stripes. Phal. ‘Taida Salu’ was produced from the cross between yellow-flower/red-spot Phal. ‘Salu Spot’ (pollen) and white-flower/red-stripe Phal. ‘Happy Beauty’ (seeds) and was used as breeding parents for 29 next-generation hybrids (Table 23.8). The white-flower/red-stripe Phal. ‘Little Gem Stripe’ was used to produce the hybrids, Phal. ‘Taida Golden Gem’ (Phal. ‘Little Gem Stripe’ × Phal. ‘Salu Peoker’) and Phal. ‘Fangmei Sweet’ (Phal. ‘Little Gem Stripe’ × Phal. ‘Dou-dii Golden Princess’) (Fig. 23.10), which have 5 next-generation hybrids and won 15 awards.

23.4.3.5 Important Hybrids with Orange Flowers

The orange-flower hybrids were produced from crosses between yellow-flower and red-flower hybrids. To get orange color, the following three events in pigment formation have to occur: first, the yellow color is stable and non-disappeared; second, the red color is distributed evenly over flowers; and third, the yellow and red colors interact together for color formation. Overall, the yellow color originated from Phal. venosa and Phal. amboinensis, and the red color came from the blood- lines of Phal. violacea and Phal. lueddemanniana. In 1961, Phal. ‘Princess Kaiulani’ was registered from the cross between Phal. violacea and Phal. amboinensis (Table 23.9). In 1974, Phal. ‘George Vasquez’ was bred from Phal. violacea and Phal. ‘Luedde-violacea’. Recently, Phal. ‘Brother Sara 598 C.-C. Hsu et al.

Gold’ (Phal. ‘Sara Lee’ × Phal. ‘Taipei Gold’), Phal. ‘Salu Peoker’ (Phal. ‘Golden Sun’ × Phal. ‘Yellow Peoker’), and Phal. ‘Sogo Lawrence’ (Phal. ‘Brother Peter Star’ × Phal. ‘Brother Sara Gold’) were important breeding parents, which contributed in breeding of 91, 86, and 66 next-generation hybrids and 351, 289, and 161 hybrids of total generations, respectively. However, most of these flowers have yellow tepals with red spots evenly distributed in whole flowers, but not like the real orange color. Moreover, Phal. ‘Surf Song’ (Phal. ‘Taipei Gold’ × Phal. ‘King Shiang’s Rose’) has pink flowers with red stripes, and one clone ‘OX Gold Orange’ has been chosen for the development of orange flowers in OX Orchids Farm (Fig. 23.11). Phal. ‘Surf

Fig. 23.11 Various phenotypes of Phalaenopsis hybrids with orange flowers. (a) Phal. ‘Brother Sara Gold’, (b) Phal. ‘Kdares Orange Lover’, (c) Phal. ‘Surf Song’, (d) Phal. ‘KS Orange’, (e) Phal. ‘Charming Fortune’, (f) Phal. ‘Fangmei A Hot’, bars=1 cm 23 Phalaenopsis 599

Song’ has 14 next-generation hybrids and gains 3 awards. Genealogy analysis showed that Phal. ‘Surf Song’ contains Phal. venosa for 25.0%, Phal. amabilis for 23.3%, Phal. rimestadiana for 22.8%, Phal. aphrodite for 10.1%, Phal. schilleriana for 9.1%, Phal. sanderiana for 4.2%, and others. The yellow color was from the bloodline of Phal. venosa, and the red color was from Phal. schilleriana and Phal. sanderiana.

23.4.3.6 Important Hybrids with Green Flowers

Green flowers are formed when the flowers contain green chloroplasts that do not degrade. The major native species for the development of green flowers were Phal. cornu-cervi var. alba, Phal. mannii var. alba, Phal. violacea, and Phal. gigantea. In 1984, Phal. ‘Gelblieber’ (Phal. amboinensis × Phal. micholitzii) was used as a major breeding patent for green-flower hybrids (Table 23.9). Its next-generation hybrids, Phal. ‘Yungho Gelb Canary’ (Phal. ‘Gelblieber’ × Phal. ‘Princess Kaiulani’), Phal. ‘Su-An Cricket’ (Phal. ‘Gelblieber’ × Phal. mannii), Phal. ‘Hannover Passion’ (Phal. ‘Gelblieber’ × Phal. mariae), and Phal. ‘Buena Jewel’ (Phal. violacea × Phal. ‘Gelblieber’), all have green tepals with white lip and are important breeding parents. From the cross between Phal. ‘Golden Sands’ and the wax-textured Phal. gigan- tea, Phal. ‘Liu Tuen-Shen’ was selected, and new progenies with better texture of flowers were obtained. Its progeny, Phal. ‘Fortune Saltzman’ (Phal. ‘Liu Tuen-­ Shen’ × Phal. ‘Barbara Freed Saltzman’) with clear light-green color, is a popular cultivar in the market (Fig. 23.12). Genealogy analysis showed that Phal. ‘Fortune Saltzman’ contains Phal. gigantea for 25.0%, Phal. amabilis for 23.3%, Phal. rime- stadiana for 20.8%, Phal. lueddemanniana for 17.2%, Phal. aphrodite for 10.1%, Phal. equestris for 3.1%, Phal. schilleriana for 0.2%, and Phal. stuartiana for 0.2%. The green flower should come from the bloodline of Phal. gigantea.

23.4.4 Harlequin (Black) Flowers

The “harlequin” flowers are called for the similarity to the face of clown with the painted black spots. Harlequin flowers contain very dark-red or dark-purple colors, which often result in very complicated color patterns (Fig. 23.13). In market, harlequin flowers are a new color in Phalaenopsis, but they were not very popular in the early stage of breeding. Two reasons for that were the harlequin color looks dirty, and the mutation rate is relative higher during micropropagation for harlequin flowers compared to other flowers. The origin of harlequin flowers can be traced back to the finding of a somaclonal mutant of Phal. ‘Golden Peoker’ “Brother” (Phal. ‘Misty Green’ × Phal. ‘Liu Tuen-­ Shen’), which was registered by Brother Orchid Nursery Co., Ltd. Its parents, Phal. ‘Misty Green’, contain green flowers with red spots, and Phal. ‘Liu Tuen-Shen’ has 600 C.-C. Hsu et al.

Fig. 23.12 Various phenotypes of Phalaenopsis hybrids with green hybrids. (a) Phal. ‘Lioulin Green Eagle’, (b) Phal. ‘Nobby’s Green Eagle’, (c) Phal. ‘Heliodor’, (d) Phal. ‘Fortune Saltzman’, (e) Phal. ‘Unimax Moonlight’, (f) Phal. ‘ARK’s Green Angel’, bars=1 cm yellow flowers with red spots. Their hybrid, Phal. ‘Golden Peoker’, contains white flowers with red spots, and “Brother” is the best selection within this hybrid. During the micropropagation of Phal. ‘Golden Peoker’ “Brother,” one somaclonal mutant was found by Ever Spring Orchids Nursery and called Phal. ‘Golden Peoker’ “Ever-­ spring (ES)”, which contain purple spots with few spots fused together. During micropropagation of Phal. ‘Golden Peoker’ “ES,” three different phenotypes can be obtained. About 40% of the micropropagated plants had the flowers similar to Phal. 23 Phalaenopsis 601

Fig. 23.13 The “harlequin” flowers of Phalaenopsis hybrids. (a) Phal. ‘Ever-spring Prince’, (b) Phal. ‘OX Black Tea’, (c) Phal. ‘Chian Xen Diamond’, (d), Phal. ‘Chian Xen Piano’, (e) Phal. ‘Lianher Focus’, (f) Phal. ‘Ebony Sweet Gem’, bars=1 cm

‘Golden Peoker’ “Brother”, 30% plants remained Phal. ‘Golden Peoker’ “ES”, and the other 30% formed a new phenotype with large-fused and dark-purple spots and were named Phal. ‘Golden Peoker’ “BL”. Both Phal. ‘Golden Peoker’ “ES” and Phal. ‘Golden Peoker’ “BL” opened a new page for Phalaenopsis breeding for har- lequin flowers. There are no native Phalaenopsis species directly related to the harlequin flow- ers. The genealogy of Phal. ‘Golden Peoker’ showed that Phal. gigantea for 25.0%, 602 C.-C. Hsu et al.

Phal. rimestadiana for 15.0%, Phal. amboinensis for 12.5%, Phal. lueddemanniana for 12.5%, Phal. amabilis for 10.2%, Phal. aphrodite for 7.4%, and others are genetically involved.

23.4.4.1 Important Hybrids with Harlequin Flowers

With the appearance of Phal. ‘Golden Peoker’ “ES,” Phal. ‘Ever-spring King’ (Phal. ‘Chih Shang’s Stripes’ × Phal. ‘Golden Peoker’) and Phal. ‘Ever-spring Light’ (Phal. ‘Ever-spring Star’ × Phal. ‘Golden Peoker’) were bred, but only 1/300 offspring contains harlequin flowers (Table 23.10). In 1999, Phal. ‘Ever-spring Prince’ (Phal. ‘Golden Peoker’ × Phal. ‘Taisuco Beauty’) was registered. By using Phal. ‘Golden Peoker’ “BL” instead of Phal. ‘Golden Peoker’ “ES,” the percent- ages of the harlequin flowers were increased from 1/300 (using “ES”) to 1/30 (using “BL”). So the generation of a relative stable harlequin-flower phenotype encour- aged breeders to continue black-flower breeding. Phal. ‘Golden Peoker’ “BL” has further been used for more harlequin-flower breeding. Phal. ‘I-Hsin Black Jack’ (Phal. ‘Golden Peoker’ × Phal. ‘Leopard Prince’), Phal. ‘Haur Jin Diamond’ (Phal. ‘Golden Peoker’ × Phal. ‘Ching-Her Buddha’), and Phal. ‘Sunrise Red Peoker’ (Phal. ‘Golden Peoker’ × Phal. ‘Kuntrarti Rarashati’) are important breeding parents and have 26, 77, and 43 next-generation hybrids and 105, 366, and 100 hybrids of total generations, respectively. Phal. ‘Ever-spring Light’ contains the most progenies of Phal. ‘Golden Peoker’ and had 26 next-generation hybrids and 899 hybrids of total generations. Its next-­ generation, Phal. ‘Ching Hua Spring’ (Phal. ‘Minho Princess’ × Phal. ‘Ever-spring Light’), is important for breeding Phal. ‘Chian Xen Pearl’ (Phal. ‘Ching Hua Spring’ × Phal. ‘Nobby’s Pink Lady’), a very important breeding parent. Phal. ‘Chian Xen Pearl’ has won 24 awards in the progenies of Phal. ‘Golden Peoker’. Other breakthrough selections which contributed significantly in the develop- ment of the assortment are Phal. ‘Ever-spring King’ and its progeny, Phal. ‘Happy Ufo’ (Phal. ‘King Shiang’s Beauty’ × Phal. ‘Ever-spring King’), and Phal. ‘Haur Jin Diamond’ and its hybrids, Phal. ‘Tai-I Yellow Bird’ (Phal. ‘Salu Peoker’ × Phal. ‘Haur Jin Diamond’) and Phal. ‘Haur Jin Princess’ (Phal. ‘Haur Jin Diamond’ × Phal. ‘Dou-dii Golden Princess’) (Table 23.10). After the appearance of Phal. ‘Golden Peoker’ mutants, Brother Orchid Nursery Co., Ltd. has used it as parent for breeding and registered for 18 G1 hybrids, includ- ing Phal. ‘Brother Purple’ and Phal. ‘Brother Peacock’. The G1 hybrids of Phal. ‘Brother Purple’, Phal. ‘Brother Precious Stones’ (Phal. ‘Brother Fancy’ × Phal. ‘Brother Purple’), Phal. ‘Brother Pirate King’ (Phal. ‘Fortune Buddha’ × Phal. ‘Brother Purple’), and Phal. ‘Brother Supersonic’ (Phal. ‘Sara Lee’ × Phal. ‘Brother Purple’), were subsequently used in further development of hybrids. 23 Phalaenopsis 603

Table 23.10 Important hybrids for the breeding of harlequin flowers from Phal. Golden Peoker and its progenies No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G0 1983 (Misty Green 208 3227 9 –c WspW Golden × Liu Peoker Tuen-Shen) Phal. G1 1992 (Chih Shang’s 74 550 9 8.0 WspR Ever-spring Stripes × King Golden Peoker) Phal. G2 2000 (King 25 157 3 6.8 –d Happy Ufo Shiang’s Beauty × Ever-spring King) Phal. G2 2002 (Ever-spring 16 21 0 – – Brother King × Sinica Love Rosa Knight) Phal. Yu G2 2001 (Ever-spring 14 27 1 7.7 WspW Pin Pearl King × Musashino) Phal. G1 1992 (Ever-spring 26 899 5 7.8 WspR Ever-spring Star × Golden Light Peoker) Phal. Ching G2 1998 (Minho 6 841 0 – WspR Hua Spring Princess × Ever-spring Light) Phal. Chian G3 2002 (Ching Hua 164 724 24 9.2 WspR Xen Pearl Spring × Nobby’s Pink Lady) Phal. G4 2005 (Chian Xen 40 110 4 10.9 PspR Fusheng Pearl × Pink Pearl Fusheng’s Purple Gem) Phal. Chian G4 2004 (Chian Xen 28 66 8 10.4 WspR Xen Pearl × Mammon Mount Beauty) Phal. Chian G4 2004 (Chian Xen 21 28 3 11.6 WspR Xen Pearl × Tinny Magpie White) Phal. Yu G4 2003 (Yu Pin Lover 20 41 5 10.0 PspR Pin Lady × Chian Xen Pearl) (continued) 604 C.-C. Hsu et al.

Table 23.10 (continued) No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G4 2004 (Chian Xen 17 55 1 10.0 PspR Fureshing Pearl × Chian Speckle Xen Sweet Valentine) Phal. Taida G3 2001 (Ching Hua 54 104 6 8.1 WspR Pearl Spring × Sogo Davis) Phal. KV G4 2006 (Salu Peoker 2 2 9 8.8 YspR Charmer × Taida Pearl) Phal. G1 1999 (Golden 46 100 13 8.1 WspR Ever-spring Peoker × Prince Taisuco Beauty) Phal. G1 1997 (Taisuco 9 9 2 10.2 WspR Ever-spring Kochdian × Fairy Golden Peoker) Phal. G1 1999 (Golden 26 105 13 7.5 WspP I-Hsin Peoker × Black Jack Leopard Prince) Phal. Haur G1 1999 (Golden 77 366 4 7.5 YspR Jin Peoker × Diamond Ching-Her Buddha) Phal. Tai-I G2 2002 (Salu Peoker 40 90 5 5.7 YspR Yellow Bird × Haur Jin Diamond) Phal. Haur G2 2002 (Haur Jin 39 91 12 7.1 YspR Jin Diamond × Princess Dou-dii Golden Princess) Phal. G2 2003 (Chingruey’s 21 29 1 – – Diamond Beauty × Beauty Haur Jin Diamond) Phal. Shin G2 2002 (Haur Jin 12 25 4 8.2 YspR Yi Diamond × Diamond Ching-Her Buddha) (continued) 23 Phalaenopsis 605

Table 23.10 (continued) No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. Yu G2 2003 (Autumn Sun 13 39 4 5.5 YspR Pin × Haur Jin Natsume Diamond) Phal. Chian G1 2002 (Golden 5 6 1 8.6 PspR Xen Peoker × Diamond Judy Valentine) Phal. G1 2003 (Golden 43 100 1 4.8 YspP Sunrise Peoker × Red Peoker Kuntrartic Rarashati) Phal. G1 1995 (Golden 97 329 0 – WspR Brother Peoker × Purple Brother Glamour) Phal. G1 1992 (Paifang’s 16 439 0 – WspR Brother Queen × Peacock Golden Peoker) Phal. G2 1998 (Brother 45 61 24 7.5 WspR Brother Fancy × Precious Brother Stones Purple) Phal. G2 1998 (Fortune 39 74 19 7.5 YspR Brother Buddha × Pirate King Brother Purple) Phal. G2 1997 (Sara Lee × 23 30 19 6.6 WspR Brother Brother Supersonic Purple) Data were collected from OrchidWiz X3.3, 2017 aFor generations, G0 is Phal. ‘Golden Peoker’. G1~G4 indicate the different generations derived from Phal. ‘Golden Peoker’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, Wsp for white with spots, Wst for white with stripes, P for pink, Psp for pink with spots, R for red, Y for yellow, Ysp for yellow with spots cNot available in OrchidWiz X3.3, 2017 dNot available in OrchidWiz X3.3, 2017 606 C.-C. Hsu et al.

23.4.4.2 Molecular Genetics for Breeding Harlequin Flowers

Harlequin or black flowers contain high amount of anthocyanin accumulations (Kuo and Wu 2011), which resulted from the high expression of anthocyanin-related genes, such as enzymes and transcription factors. In blood oranges (Citrus sinensis), a Copia-like retrotransposon controls the high expression of a nearby MYB transcription factor, Ruby, and results a large amount of anthocyanin production in the fruits (Butelli et al. 2012). A similar situation has also been found in cauliflower, where a Harbinger DNA transposon was inserted in the regulatory region of Purple (Pr) gene, encoding for a R2R3-MYB transcription factor, and resulted in the upregulation of Pr and intense purple color in curds (Chiu et al. 2010). It is possible that the harlequin flowers in Phalaenopsis are caused by transposon insertions in the regulatory regions of a R2R3-MYB transcription factor.

23.4.5 Purple-to-Blue Flowers

Although there are 34,112 hybrids of Phalaenopsis registered in RHS, the hybrids with flowers containing blue or blue-to-purple color are very rare. Moreover, the “blue” flowers in Phalaenopsis are indigo, purple-to-blue, or lavender colors, but not the sky blue or so-called true blue. In nature, most species of Phalaenopsis do not have blue flowers; however, a few varieties have been found with purple-to-blue colors, so the dreaming blue flowers may not be a dream.

23.4.5.1 Native Species Contributing to Purple-to-Blue Hybrids

Several varieties of native Phalaenopsis species have been found with purple-to-­ blue flowers, includingPhal. pulcherrima var. coerulea, Phal. equestris var. coeru- lea, Phal. bellina, and Phal. violacea “Indigo” (Fig. 23.14). Phal. pulcherrima contains various flower colors, from white, dark red, to purple-to-blue, and Phal. pulcherrima var. coerulea contains purple-to-blue tepals and dark-purple lip. Phal. equestris contains various flower colors, including white, pink, and red, and Phal. equestris var. coerulea has the white tepals with purple-to-blue lip. Phal. bellina contains yellow and purple flowers, and Phal. bellina “Indigo” has purple-to-blue flowers, although its offspring has returned to purple flowers. Phal. violacea contains dark-purple flowers, and Phal. violacea “Indigo” has purple-to-blue flowers. Another genus of Orchidaceae, Vanda, has closer purple-to-blue color than Phalaenopsis flowers, so the introduction of the bloodline of Vanda into Phalaenopsis will be an alternative way for breeding purple-to-blue flowers. 23 Phalaenopsis 607

Fig. 23.14 Flowers of purple-to-blue Phalaenopsis species. (a) Phal. equestris, (b) Phal. viola- cea, (c) Phal. bellina, (d) Phal. ‘Kenneth Schubert’, (e) Phal. Purple Martin, bars=1 cm

23.4.5.2 Important Hybrids with Purple-to-Blue Flowers

Phal. ‘Kenneth Schubert’ (Phal. pulcherrima × Phal. violacea) was the first regis- tered hybrid with purple-to-blue flowers and used as a major breeding parent (Fig. 23.14). Its offspring, Phal. ‘Purple Martin’ (Phal. ‘Kenneth Schubert’ × Phal. violacea) and Phal. ‘Little Blue Bird’ (Phal. ‘Kenneth Schubert’ × Phal. pulcher- rima), were from the backcross of Phal. ‘Kenneth Schubert’ to its parents. So far, Phal. ‘Purple Martin’ has the closest blue-colored flowers in Phalaenopsis. Recently, with the finding of the new varieties from native species, the breeding for purple-to-blue and blue flowers has got a new impulse. 608 C.-C. Hsu et al.

23.4.5.3 Molecular Genetics for Breeding Purple-to-Blue Flowers

The major pigment compounds for flower colors are anthocyanins and carotenoids. Anthocyanins contribute to the orange, red, to blue colors, while carotenoids are responsive to the yellow color. There are three kinds of anthocyanins present in plants and related to different colors, pelargonidin, cyanidin, and delphinidin for orange, red, and blue colors, respectively. The transition from pelargonidin to cyani- din or cyanidin to delphinidin needs the enzyme activities of flavonoid-3′- hydroxylase (F3’H) or flavonoid-3′5′-hydroxylase (F3′5′H), respectively (Sasaki and Nakayama 2015). For example, salvia does not have F3’H and F3′5′H and produces pelargonidin-derived anthocyanin to show orange-to-red color (Tanaka and Ohmiya 2008). Rose has F3′H without F3′5′H, so it creates cyanidin-kind anthocyanin for red color (Tanaka and Ohmiya 2008). Lavender contains both F3′H and F3′5′H and produces delphinidin-type anthocyanin and results in blue color (Tanaka and Ohmiya 2008). Therefore, if breeders want to have blue flowers for Phalaenopsis orchids, it needs to have functional F3′5′H expressed in the flowers. Previous studies showed that the gene encoding F3′5′H enzyme has been cloned from black-flower, red-flower Phalaenopsis hybrids, and its function has been investigated by transient expression assay, although the transformed flowers did not show blue color (Su and Hsu 2003; Wang et al. 2006). In addition, the pH value within the vacuole, where the anthocyanin is present, has been reported to affect the purple-to-blue color, meaning that the flower containing cyanidin looked bluer when pH value increased in the vacuole and became redder under the low pH value (Griesbach 2005).

23.5 Phalaenopsis Breeding for Differential Morphology

Besides color, flower morphology is one of the most important features for breeding Phalaenopsis. Breeding for smaller flower sizes and different flower shapes are becoming more important during the last decades.

23.5.1 Small and Medium Flowers

Most Phalaenopsis breeding has emphasized on large flowers (>10 cm) with vari- ous colors; see Sects. 23.4.1 and 23.4.2. Following the saturation of the breeding for large flowers, a new breeding direction for small- and medium-sized flowers has emerged recently. Typically the plants are around 20 cm in height and the flowers are 3~7 cm in width. The small- and medium-sized flowers are economically profitable as more plants can be produced per m2 of greenhouse space and plants are 23 Phalaenopsis 609

Fig. 23.15 Flowers of small Phalaenopsis species, Phal. equestris, with (a) white sepal/petal and white lip, (b) white sepal/petal and yellow lip, (c) red sepal/petal and red lip, (d) red sepal/petal and orange lip, bars=1 cm easy to transport. In addition, the small- and medium-flower hybrids are suitable as gifts or are used to decorate office tables or writing desks, which means that these hybrids have a different market from the large-flower hybrids.

23.5.1.1 Native Species Contributing to Small and Medium Flowers

The major native species for small flowers are Phal. equestris, Phal. pulcherrima, and Phal. lindenii. Phal. equestris that contain many flowers in one inflorescence, and the flowers are 2.5 cm in width in various colors, from white, pink, red, to purple-red (Fig. 23.15). The interesting genetic features of Phal. equestris for breeding programs are compact plants and small flowers, more floral stalks with many flowers, and various flower colors (Hsu and Chen 2015). Phal. pulcherrima has flowers with 2~3 cm in width and various colors and attributes to the develop- ment of dark-red and dark-purple colors. Flowers of Phal. lindenii are about 3 cm in width, and this species is the major resource to introduce the red-stripped feature. 610 C.-C. Hsu et al.

Table 23.11 Important hybrids for the breeding of small and medium flowers from Phal. ‘Cassandra’ and its progenies No. of Total Year next-­ no. of Natural registered generation hybrids spread Hybrid name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G0 1896 (equestris × 222 3305 25 4.5 PR Cassandra stuartiana) Phal. G1 1982 (Cassandra × 197 615 8 5.5 WW Timothy aphrodite) Christopher Phal. Liu’s G2 2004 (Rainbow Chip 46 71 3 3.6 WstR Twilight × Timothy Rainbow Christopher) Phal. Liu’s G2 2000 (lindenii × 30 76 1 4.0 WstR Bright Timothy Ruby Christopher) Phal. Rong G2 1996 (Timothy 29 86 11 5.2 WY Guan Amah Christopher × amabilis) Phal. Tying G2 2008 (Chian Xen 3 3 17 7.1 WspR Shin Pearl × Fantastic Timothy World Christopher) Phal. G1 1990 (Terilyn 159 1221 22 5.5 PR Carmela’s Fujitake × Pixie Cassandra) Phal. G2 1992 (Carmela’s 131 710 25 3.8 RR Zuma’s Pixie × Pixie equestris) Phal. Ho’s G2 1993 (Be Glad × 42 89 1 4.0 WstR Little Carmela’s Caroline Pixie) Phal. G2 1994 (Sun-Chen 41 165 6 6.1 RR Taisuco Beauty × Pixie Carmela’s Pixie) Phal. Sogo G2 1996 (Happy Lip × 29 46 1 5.6 WR Mini Dog Carmela’s Pixie) Phal. Super G2 1996 (Carmela’s 18 28 0 –c –d Pixie Pixie × Zuma’s Pixie) Phal. Be G1 1978 (Swiss Miss × 131 1351 18 4.9 WR Glad Cassandra) Phal. Be G2 1989 (Be Glad × 75 351 19 4.0 PR Tris equestris) (continued) 23 Phalaenopsis 611

Table 23.11 (continued) No. of Total Year next-­ no. of Natural registered generation hybrids spread Hybrid name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G2 1998 (Be Glad × 51 81 14 5.8 WspW Nobby’s Rothschildiana) Amy Phal. Ho’s G2 1993 (Be Glad × 42 89 1 4.0 WstR Little Carmela’s Caroline Pixie) Phal. Ho’s G2 1990 (Be Glad × 40 145 13 6.2 WspR Amaglad amabilis) Phal. Liu’s G2 1995 (Be Glad × 38 67 0 – WspY Rainbow lindenii) Phal. Glad G2 1985 (Be Glad × 34 193 2 8.1 WR Melinda Melinda Nan) Phal. Pixie G5 1997 (pulcherrima × 5 5 21 3.8 RR Star Joyful) Phal. G1 2008 (Cassandra × 38 42 13 6.4 WspW Nankung’s Nankung’s 4.55 PM Beauty Girl) Phal. G1 1970 (Cassandra × 37 96 0 PP Caribbean Mambo) Sunset Phal. Purple –e 1963 (pulcherrima × 29 41 32 3.2 RR Gem equestris) Phal. – 1956 (amabilis × 5 6 1 – WW Eduardo pulcherrima) Quisumbing Data were collected from OrchidWiz X3.3, 2017 aFor generations. G0 is Phal. ‘Cassandra’. G1~G5 indicate the different generations derived from Phal. ‘Cassandra’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, Wsp for white with spots, Wst for white with stripes, P for pink, R for red, Y for yellow cNot available in OrchidWiz X3.3, 2017 dNot available in OrchidWiz X3.3, 2017 eNot the progenies of G0, Phal. ‘Cassandra’

23.5.1.2 Important Hybrids with Small and Medium Flowers

The major breeding parent for small and medium hybrids is Phal. ‘Cassandra’ (Phal. equestris × Phal. stuartiana). This hybrid was registered in 1896. It has flowers of 3~5 cm in pink colors with red spots. In history this hybrid had contributed to 222 G1 generations and 3305 progenies in total generations (Table 23.11; Fig. 23.16). Its off- spring, Phal. ‘Timothy Christopher’ (Phal. ‘Cassandra’ × Phal. aphrodite), Phal. ‘Carmela’s Pixie’ (Phal. ‘Terilyn Fujitake’ × Phal. ‘Cassandra’), and Phal. ‘Be Glad’ (Phal. ‘Swiss Miss’ × Phal. ‘Cassandra’), have the most next-generation offspring. 612 C.-C. Hsu et al.

Fig. 23.16 Various phenotypes of Phalaenopsis hybrids with small flowers. (a) Phal. ‘Cassandra’, (b) Phal. ‘Liu’s Hua Lien Red-Carpet’, (c) Phal. ‘Taida Little Vivien’, (d) Phal. ‘Brother Spring Dancer’, bars=1 cm

Flowers of Phal. ‘Timothy Christopher’ are 4.0~6.2 cm in width and have a white color. Its G2 hybrid, Phal. ‘Liu’s Twilight Rainbow’ (Phal. ‘Rainbow Chip’ × Phal. ‘Timothy Christopher’), is the hybrid with the most flowers in all registered Phalaenopsis hybrids. In total 248 flowers were counted in one plant. Comparing all the progenies resulting from Phal. ‘Timothy Christopher’, Phal. ‘Tying Shin Fantastic World’ (Phal. ‘Chian Xen Pearl’ × Phal. ‘Timothy Christopher’) won the most awards with 17 awards. Phal. ‘Carmela’s Pixie’ has 5 cm flowers with various colors from pink, purple-­ red, to dark-red. Its next-generation hybrid, Phal. ‘Zuma’s Pixie’ (Phal. ‘Carmela’s Pixie’ × Phal. equestris), has still smaller flowers (3.2~4.5 cm) and colors from purple-red to dark-purple with a dark-red lip. Among the different generated prog- enies of Phal. ‘Carmela’s Pixie’, Phal. ‘Zuma’s Pixie’ was the most successful one. Phal. ‘Be Glad’ flowers are 3.8~5.3 cm in width and white colored with blushed pink on the bases and a dark-rose lip. Its progeny, Phal. ‘Be Tris’ (Phal. ‘Be Glad’ × Phal. equestris), has flowers of 3.0~4.6 cm in width and was frequently used in breeding. Among the progenies of Phal. ‘Be Glad’, Phal. ‘Pixie Star’ (Phal. pul- cherrima × Phal. ‘Joyful’) and Phal. ‘Be Tris’ won the most awards (Table 23.11). The other primary hybrids from the cross between two native species, such as Phal. ‘Purple Gem’ (Phal. pulcherrima × Phal. equestris) and Phal. ‘Eduardo 23 Phalaenopsis 613

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0 Generations 2~2.93~3.9 4~4.95~5.9 6~6.97~7.9 8~8.99~9.9 10~10.911~11.9 12~12.913~13.9 14~14.915~15.9 Flower sizes (cm) 13 001 00000000000 12 0 0 0 21010000000 11 00100000000000 10 00021010000000 9 02165622111000 8 011916 36 25 91618 9200 7 01212 38 75 59 49 44 29 15 830 6 02317 58 80 93 87 53 35 28 700 5 01313 57 77 83 79 84 54 23 10 10 4 00014 34 67 86 101 65 34 13 100 3 021220 38 57 48 30 17 1100 2 0872535372420114 0001 1 115 49 64 63 26 12 6310000 018 47 58 36 912000000 614 39 22 19 110000000

Fig. 23.17 Relationships of the flower sizes of the progenies of Phal. equestris from various gen- erations, and data were from Orchidwiz X3.3, 2017

Quisumbing’ (Phal. amabilis × Phal. pulcherrima), were also used in breeding for small- and medium-sized flowers. Phal. ‘Purple Gem’ contains flowers with 2.5~3.9 cm in width and various colors from dark pink to dark purple, while Phal. ‘Eduardo Quisumbing’ is characterized by white flowers and a straight floral stalk without branched. In order to investigate how Phal. equestris contributed in the development of small and medium hybrids, the flower sizes in the different progenies ofPhal. equestris were analyzed. In total 2774 of the 21,805 progenies that were recorded in RHS (OrchidWiz 2017) to have Phal. equestris background were analyzed (Fig. 23.17). The data showed that more progenies from the first to the third generations have flower sizes between 4 and 6 cm, while more progenies from the fifth to the tenth generations have flowers with 6–11 cm (Fig. 23.17). These results show that the hybrids derived from the first generations after crossing withPhal. equestris, which have higher percentages of the bloodline from Phal. equestris, own the features of small and medium flowers.

23.5.2 Peloric Flowers

Peloric flowers have two petals becoming lip-like morphology and seem to contain three sepals, three lips, and one column (Fig. 23.18). The name of “peloric flower” can be traced to Charles Darwin in his The Variation of Animals and Plants under Domestication for snapdragon (Antirrhinum) (Darwin 1868) and referred to the 614 C.-C. Hsu et al.

Fig. 23.18 Various phenotypes of Phalaenopsis hybrids with peloric flowers. (a) Normal type and (b, c) peloric type of Phal. equestris, (d) Phal. ‘Liu’s Pale Micholitz’, (e) Phal. ‘Fuller’s Miss’, bars=1 cm aberration in flowers from normal zygomorphy to mutant actinomorphy. Most peloric flowers occurred as mutation and are given up by orchid farmers, but few peloric flowers with slight changed lip-like petals are chosen to be new hybrids. Peloric flowers contain two petals changing to lip-like shape. The molecular mechanism for the lip development is an important and interesting question for Orchidaceae. A floral organ identity gene, B-class PeMADS4, has been reported as a regulator for determining the lip development (Tsai et al. 2004). In addition, another group, Agamous-like 6 (AGL6) genes, interacts with B-class genes to regulate the lip or tepal development together in Phalaenopsis flowers (Hsu et al. 2015b).

23.5.3 Bigfoot Flowers

In opposite to the peloric flowers, the Bigfoot flowers have the lip becoming petal-­ like morphology, so these flowers seem to have three sepals, three petals, and one column that make the flowers nearly round shape (Fig. 23.19). Actually, the Bigfoot flowers are one kind of peloric flowers but were named after the first breeding -par ent, Phal. ‘World Class’ “Bigfoot” (Phal. ‘Mae Hitch’ × Phal. ‘Kathy Sagaert’). This hybrid had the Bigfoot phenotype in all 11 flowers on a floral stalk and was 23 Phalaenopsis 615

Fig. 23.19 Flowers of Bigfoot hybrids. (a) Phal. ‘United White Bear’, (b) Phal. ‘Fangmei Dream Wedding’, (c) Phal. ‘Fuller’s 3580’, (d) Phal. ‘Taisuco Sunstone’, (e) Phal. ‘Lioulin Lovely Lip’, (f) ‘Fuller’s O-Plus’, bars=1 cm selected by Carmela Orchids (HI, USA) in 1991. Phal. ‘World Class’ “Bigfoot” was the only individual containing Bigfoot phenotype in the G1 population of the cross of Phal. ‘Mae Hitch’ to Phal. ‘Kathy Sagaert’, and its parents showed no Bigfoot phenotype. So the Bigfoot trait seems to result from gene mutation, but not due to genetic inheritance. So the breeding of Bigfoot flowers is very recent, and their sales in Phalaenopsis market started during the last decade. The Bigfoot phenotype is variable with the slightly to severely changed petal-like lip, and the flower colors are white, pink, red, and white with red spots and stripes. 616 C.-C. Hsu et al.

There were no direct native Phalaenopsis species contributing to Bigfoot flow- ers. Genealogy analysis showed that Phal. ‘World Class’ “Bigfoot” contains Phal. rimestadiana for 26.40%, Phal. amabilis for 25.30%, Phal. equestris for 13.60%, Phal. schilleriana for 12.3%, Phal. aphrodite for 11.40%, Phal. sanderiana for 5.80%, Phal. stuartiana for 4.00%, and Phal. lueddemanniana for 1.20%.

23.5.3.1 Important Hybrids with Bigfoot Flowers

Phal. ‘World Class’ “Bigfoot” is the beginning of the so-called Bigfoot flowers and used as breeding parents for 44 next-generation hybrids and 355 hybrids of total generations (Table 23.12). The breeding for Bigfoot flowers was not widely started, because few breeders own the plants of Phal. ‘World Class’ “Bigfoot” or its off- spring. Until 2011, most hybrids were registered due to the occurrence and spread- ing of Phal. ‘Yu Pin Easter Island’ (Phal. ‘Sogo Yukidian’ × Phal. ‘World Class’) in 2009 and Phal. ‘Yu Pin Fireworks’ (Phal. ‘Sogo Dove’ × Phal. ‘World Class’) in 2010. From 2012 to 2016, 38 to 63 hybrids were registered each year (Fig. 23.20), and the breeding of Bigfoot flowers has started extensively. Phal. ‘Yu Pin Easter Island’ and Phal. ‘Yu Pin Fireworks’ were registered in RHS by Neng-I Chang (Yu Pin Biological Technology Co., Ltd., Taiwan) and were sold as normal Phalaenopsis flowers. This fact increased the spreading of the breed- ing for Bigfoot flowers. As the G1 offspring of Phal. ‘World Class’ “Bigfoot,” Phal. ‘Yu Pin Easter Island’ and Phal. ‘Yu Pin Fireworks’ had 64 and 79 next-generation hybrids and 112 and 161 hybrids of total generations, respectively (Table 23.12). Phal. ‘Yu Pin Easter Island’ has 7~14 cm flowers and white color with dark-pink stripes, while Phal. ‘Yu Pin Fireworks’ contains flowers of 8~13 cm in width and white color with pink stripes. By using Phal. ‘Yu Pin Easter Island’ and Phal. ‘Yu Pin Fireworks’ as breeding parents, Fuller’s Orchid Nursery registered 13 and 25 next-generation hybrids, respectively; Tying Shin Orchid registered 1 and 13 hybrids, respectively; and Huang Gao Ming registered 9 and 5 hybrids, respectively. Among them, Phal. ‘Fuller’s AD-Plus’ (Phal. amabilis × Phal. ‘Yu Pin Easter Island’) from Fuller’s Orch., Phal. ‘Tying Shin World Class’ (Phal. ‘Sogo Vivien’ × Phal. ‘Yu Pin Fireworks’) from Tying Shin Orch., Phal. ‘Lioulin Pretty Lip’ (Phal. ‘Yu Pin Fireworks’ × Phal. ‘Chian Xen Mammon’) from Mr. Huang Gao Min, and Phal. ‘Fuller’s E-Plus’ (Phal. ‘Yu Pin Fireworks’ × Phal. ‘Fuller’s Milk’) from Fuller’s Orch. were further most frequently used in breeding (Table 23.12).

23.6 Phalaenopsis Breeding for Scented Flowers

Scent is the major feature for plants, especially for the ornamental plants. Scent is a complex of fragrant compounds that are volatile and is used to attract the pollinators for helping plant pollination in nature. 23 Phalaenopsis 617

Table 23.12 Important hybrids for the breeding of Bigfoot flowers from Phal. ‘World Class’ and its progenies No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G0 1990 (Mae Hitch × 44 355 3 9.1 WstW World Kathy Sagaert) Class Phal. Yu G1 2009 (Sogo 64 112 11 10.6 WstP Pin Easter Yukidian × Island World Class) Phal. G2 2000 (amabilis × 7 76 1 7.9 WW Fuller’s Yu Pin Easter AD-Plus Island) Phal. G2 2012 (Yu Pin 5 5 0 –c –d Fuller’s Easter Island D-Plus × Fuller’s Purple Queen) Phal. G2 2012 (Yu Pin 5 5 0 – – Lioulin Easter Island Pure Lip × Sogo Yukidian) Phal. G2 2012 (Yu Pin 4 4 0 – – Fuller’s Easter Island F-Plus × Fuller’s Cow) Phal. G2 2011 (Leopard 4 4 0 – – Yushan Prince × Yu Mongo Pin Easter Island) Phal. Yu G1 2010 (Sogo Dove × 79 161 8 9.7 WstP Pin World Class) Fireworks Phal. G2 2011 (Sogo Vivien 15 16 3 6.0 WstW Tying Shin × Yu Pin World Fireworks) Class Phal. G2 2012 (Yu Pin 10 10 0 – – Lioulin Fireworks × Pretty Lip Chian Xen Mammon) Phal. G2 2012 (Yu Pin 9 9 0 – – Fuller’s Fireworks × E-Plus Fuller’s Milk) Phal. G2 2011 (Champion 7 7 0 – – Tying Shin Lightning × Red Yu Pin Emperor Fireworks) (continued) 618 C.-C. Hsu et al.

Table 23.12 (continued) No. of Total Year next-­ no. of Natural Hybrid registered generation hybrids spread name Gena on RHS Parents hybrids derived Award (cm) Colorb Phal. G2 2012 (Yu Pin 6 7 1 7.8 WstW Lioulin Fireworks × Lovely Lip Chian Xen Piano Phal. G1 2005 (World Class 7 8 2 7.5 WP Sasquatch × amabilis) Phal. G1 1994 (Paul Tatar × 6 9 0 – – Tropical World Class) Stripes Phal. G1 1996 (Carmela’s 4 5 1 6.4 PstR Jordan’s Pixie × World Pixie Class) Phal. G1 1992 (Music × 0 0 12 10.2 PstP Krull’s World Class) Butterfly Phal. G1 2008 (Taisuco 0 0 7 8.2 PstP Dendi’s Happybeauty Yeti × World Class) Data were collected from OrchidWiz X3.3, 2017 aFor generations. G0 is Phal. World Class. G1~G2 indicate the different generations derived from Phal. ‘World Class’ bThe colors for cultivar flowers. First word is the color for tepals, and second word is the color for lip. W indicates white, Wsp for white with spots, Wst for white with stripes, P for pink cNot available in OrchidWiz X3.3, 2017 dNot available in OrchidWiz X3.3, 2017

Fig. 23.20 Hybrid numbers of Phal. ‘World Class’ from 1994 to 2017 23 Phalaenopsis 619

In Phalaenopsis most breeding was focused on choosing big flowers and various colors. Few breeding was focused on selection of hybrids with scent. Furthermore most hybrids lost the scented fragrance in their next-generation offspring. The major bottleneck is that scent is not a dominant or stable trait to inherit to next generation, meaning that the scent feature in most offspring is lost or faded. In addition, most native species with strong scents are with small flowers of star shape and a limited flower number. These features of the native scented species are far from the prefer- ence of most hybrids with large rounded flowers in the market. To breed a hybrid with large and rounded flowers, the native species with scents should cross to large flowers for several generations, but during these generations, scent might be lost.

23.6.1 Native Species Contributing to Scented Flower Hybrids

Several native species of Phalaenopsis have scent feature, but only few of them contain strong fragrance and are used as breeding parents. The major native species with strong scents are Phal. amboinensis, Phal. bellina, Phal. lueddemanniana, Phal. venosa, and Phal. violacea (Table 23.13; Fig. 23.21) (Hsiao et al. 2008; Yeh et al. 2012). These species contain various kinds of volatile compounds, such as Phal. bellina having monoterpene, linalool, and geraniol as the major fragrant compounds (Hsiao et al. 2006, 2008). However, most native species with strong scents have small flowers with star shape and need to be crossed with the hybrid with large and rounded flowers. The other native species with weak to strong scents are listed in Table 23.13, but their scents are too weak or fragrant compounds are so different and hardly to be used as breeding parents for scented flowers.

23.6.2 Important Hybrids with Scented Flowers

Yeh et al. (2012) analyzed the percentages of fragrance present among Phalaenopsis hybrids (Table 23.14). The cross of scented Phal. schilleriana to scentless Phal. ‘Ho’s Little Caroline’ showed 52.7% of G1 hybrids contained scent feature, but the cross of scented to scentless Phal. ‘Sogo Berry’, Phal. ‘Be Glad’, and Phal. ‘Jiaho Cherry’ resulted in 10.2–20.0% of G1 hybrids with fragrance. In addition, the cross of scented Phal. schilleriana to scentless Phal. ‘Rong Guan Mary’ showed 0% of G1 hybrids contained scent feature. When analyzing the genealogy of these male parent, there were 1.6%, 6.1%, 0%, 1.3%, and 17.4% of the bloodline of Phal. schil- leriana in Phal. ‘Ho’s Little Caroline’, Phal. ‘Sogo Berry’, Phal. ‘Be Glad’, Phal. ‘Jiaho Cherry’, and Phal. ‘Rong Guan Mary’, respectively. All these results suggest that the scent feature is either not dependent on the female parent or relied on the percentage of the bloodline of Phal. schilleriana in the progenies. Using scented Phal. ‘Kung’s Roth-Fairy’ in crosses with scented Phal. ‘Lung-An Mist Pink’ resulted in the 57.1% of G1 hybrids with fragrance, and the cross with 620 C.-C. Hsu et al.

Table 23.13 The scents trait Scents trait for native scented species Phal. amabilis No or weak Phal. amboinensis Strong Phal. bellina Strong Phal. corningiana Strong Phal. cornu-cervi Medium Phal. equestris Weak to medium Phal. fasciata Medium Phal. fimbriata Weak Phal. fuscata Medium Phal. gigantea Weak to medium Phal. hieroglyphica Weak Phal. javanica Medium Phal. kunstleri Weak Phal. lobbii Medium to strong Phal. lueddemanniana Medium to strong Phal. mannii Weak Phal. mariae Weak Phal. modesta Strong Phal. parishii Medium Phal. pulchra Medium Phal. reichenbachiana Medium Phal. schilleriana Medium to strong Phal. stuartiana Weak to medium Phal. sumatrana Medium Phal. tetraspis Medium Phal. venosa Strong Phal. violacea Strong Phal. viridis Weak Phal. wilsonii Medium Data were modified from Hsiao et al. (2008) and Yeh et al. (2012)

Fig. 23.21 Flowers of scented Phalaenopsis species. (a) Phal. bellina, (b) Phal. violacea, bars=1 cm 23 Phalaenopsis 621

Table 23.14 Percentages of scented offspring of Phalaenopsis hybrids Scented ratio Parents Scented Scentless (%) Phal. schilleriana “HDAIS#2” × Phal. Ho’s Little Caroline 29 26 52.7 “Ho’s Little Zebra” Phal. schilleriana “HDAIS#2” × Phal. Sogo Berry 13 52 20.0 Phal. schilleriana “HDAIS#3” × Phal. Be Glad “La Flora” 38 333 10.2 Phal. schilleriana “HDAIS#3” × Phal. Jiaho Cherry 8 44 15.4 “KF#1” Phal. schilleriana “HDAIS#5” × Phal. Rong Guan Mary 0 7 0.0 Phal. Kung’s Roth-Fairy × Phal. Be Glad “La Flora” 0 49 0.0 Phal. Kung’s Roth-Fairy × Phal. Lung-An Mist Pink 4 3 57.1 Phal. Kung’s Roth-Fairy × Phal. Timothy Christopher 3 4 42.9 “KF#18” Phal. Kung’s Roth-Fairy × Phal. Yu Pin Summer 0 65 0.0 This table was modified from Yeh et al. (2012)

scentless Phal. ‘Timothy Christopher’ showed 42.9% offspring having scents. However, scented Phal. ‘Kung’s Roth-Fairy’ crossed with scentless Phal. ‘Be Glad’ or Phal. ‘Yu Pin Summer’ resulted in 0% offspring with scents. These results sug- gest that a cross between two scented parents has more than 50% G1 hybrids con- taining scents, while in crosses between scented and scentless parents results are varying a lot.

23.6.3 Molecular Genetics for Breeding Scented Flowers

The molecular mechanism of scent flowers has been studied in Phal. bellina (Hsiao et al. 2008). The major scent compounds of Phal. bellina are linalool and geraniol, both belong to monoterpene. Based on the comparative transcriptome analysis, the gene encoding geranyl diphosphate synthase (GDPS) has been isolated and named PbGDPS (Hsiao et al. 2008). The expression of PbGDPS is correlated to the maximal emission of monoterpenes on day 5 post-anthesis, and the enzymatic activity is proved for catalyzing the formation of geranyl diphosphate (GDP), the precursor for the production of linalool and geraniol (Hsiao et al. 2008). Although PbGDPS is important for the scent production in Phal. bellina, there are still many questions that remain to be solved for the scent inheritance, for instance, whether the scent can be produced in scentless white-flower Phal. ‘Sogo Yukidian’ “V3” with genetic transformation of PbGDPS. Other native species with strong scents, such as Phal venosa and Phal. amboinensis, contain various fragrant compounds, but not monoterpenes. Therefore, the biosynthetic and regulatory mechanisms in various scented flowers need to be further investigated. 622 C.-C. Hsu et al.

23.7 Genome Size Variation in Species Affecting Phalaenopsis Breeding

23.7.1 Cytogenetic Study of Phalaenopsis Species

For the breeding of new varieties in Phalaenopsis, native species are often used as one of the parents to introduce specific traits, such asPhal. bellina for scents or Phal. amboinensis for yellow flowers. However, during interspecific hybridization, the sterility problem has always been encountered by breeders either in the parental crosses or the subsequent crosses of the progenies (Chuang et al. 2008). Two major causes of sterility can be identified: the difference in chromosome sizes and differences in ploidy levels among species and varieties (Arends 1970; Griesbach 1985; Chuang et al. 2008). Although all diploid species have the same number of 38 chromosomes (2n = 2x = 38) (Christenson 2001), karyotypes of 9 species showed that their chromosomes vary in size and centromere position. Among them, the chromosomes are small in Phal. aphrodite, Phal. equestris, and Phal. stuartiana (<2 μm), small and medium in Phal. cornu-cervi and Phal. lueddemanniana (<2 μm and 2.0~2.5 μm), and small, medium, and large in Phal. venosa, Phal. amboinensis, Phal. mannii, and Phal. violacea (<2 μm, 2.0~2.5 μm, and >2.5 μm) (Table 23.15) (Shindo and Kamemoto 1963; Kao et al. 2001). The commercial varieties/hybrids are usually tetraploid, while the native species are diploid (Griesbach 1985; Chuang et al. 2008). In order to have an efficient breeding program, there is a need for the information of the genome sizes and ploidy levels of the breeding materials, including the native species and the parental stocks of the Phalaenopsis. This can be accomplished by using flow cytometry with the staining of propidium iodide (PI) and 4′,6-diamidino-2-phenylindol (DAPI) (Lin et al. 2001; Chen et al. 2013b; Chen et al. 2014). From these studies in Phalaenopsis species, the variation in 2C-values of 50 species was determined and values ranged

Table 23.15 Karyotypes of somatic metaphase chromosomes in nine Phalaenopsis species Chromosome size (μm) Small Medium Large Species 2n <2.0 2.0–2.5 >2.5 Nuclear DNA content (pg 2C−1) Phal. aphrodite 38 38 0 0 2.80 ± 0.06 Phal. equestris 38 38 0 0 3.37 ± 0.05 Phal. stuartiana 38 38 0 0 3.13 ± 0.07 Phal. cornu-cervi 38 34 4 0 6.44 ± 0.16 Phal. lueddemanniana 38 28 10 0 6.49 ± 0.22 Phal. venosa 38 12 4 22 9.52 ± 0.27 Phal. amboinensis 38 8 4 26 14.36 ± 0.19 Phal. violacea 38 6 6 26 15.03 ± 0.21 Phal. mannii 38 0 6 32 13.50 ± 0.12 This table was modified from Kao et al. (2001) and Lin et al. (2001) 23 Phalaenopsis 623 from 2.80 pg in Phal. aphrodite to 15.03 pg in Phal. violacea. The measured genome size correlated well to the chromosome sizes (Table 23.15). Phal. aphrodite, Phal. equestris, and Phal. stuartiana with small chromosomes have mean 2C-values ranging from 2.8 to 3.37 pg, while Phal. cornu-cervi and Phal. lueddemanniana with small and medium have mean 2C-values from 6.44 to 6.49 pg. In addition, Phal. venosa, Phal. amboinensis, Phal. mannii, and Phal. violacea with small, medium, and large chromosomes had 2C-values from 9.52 to 14.36 pg. Overall, the chromosome sizes and genome contents of native species are highly variable and result in the sterility problem for Phalaenopsis breeding. With the cytogenetic studies, it is easy to choose more suitable species or hybrids as breeding parents to enhance the efficiency of Phalaenopsis breeding. The induction of polyploids in Phalaenopsis species provides a good strategy to enable efficient hybridization with tetraploid hybrids (see next section).

23.7.2 Induction of Polyploidy in Phalaenopsis

Polyploidy plays an important role in the breeding of many crops as well as horti- cultural plants (Sattler et al. 2016) and is also useful for the improvement of Phalaenopsis. Most of the commercial hybrids of Phalaenopsis are tetraploids, while most of the native species are diploid. Thus, a hybridization barrier between hybrids and species limits the source of germplasm available for the breeding programs. Hence a simple technique to scale up the ploidy level of diploid species is needed. Due to the occurrence of endopolyploid cells in the tissues of protocorms and protocorm-like bodies (PLBs) of Phalaenopsis, there is a chance to regenerate polyploid plants from these cells. For induction of polyploidy, a group of protocorms or PLBs are cleaved at a point of half length, and the portion with growing point is used to grow the next-­ generation PLBs in T2 medium (Hyponex No. 1 3.5 g/L, tryptone 1 g/L, citric acid 0.1 g/L, activated charcoal 1 g/L, sucrose 20 g/L, potato 20 g/L, banana 25 g/L, and agar 7.5 g/L) for 2~3 months (Chen et al. 2013a). This process is repeated several times until next-generation PLB are identified with doubled ploidy level. These PLBs are then selected and regenerated to plantlets for 5 months (Chen et al. 2013a). This is a simple, effective, and reliable technique to produce large amount of polyploid plants in Phalaenopsis orchids. It might have a great impact on the new variety development of the orchid in the future.

23.8 Conclusions and Perspectives

Phalaenopsis is the most popular cultivated orchid with a unique and beautiful flower morphology. Up to May 2017, there were 34,112 hybrids ofPhalaenopsis registered in RHS. The large flowers with white and various colors are most popular 624 C.-C. Hsu et al. in Phalaenopsis market. The colorful flowers include the pink to red and purple, yellow to green and orange, black, and various flower colors with red lip. In addition, the white, pink, or yellow flowers with various pigmentation patterning, such as red stripes and spots, have been one of the major breeding directions recently. Flowers with changed morphology were rarely found. However the peloric and Bigfoot flowers formed a new trend of Phalaenopsis breeding. In addition, breeding a scented commercial hybrid with many large and rounded flowers is a challenge for the breeders. Possibly increased knowledge of molecular biology and the use of genetic transformation can enhance this process.

References

Arends JC (1970) Cytological observations on genome homology in eight interspecific hybrids of Phalaenopsis. Gentica 41:88–100 Borevitz JO, Xia Y, Blount J, Dixon RA, Lamb C (2000) Activation tagging identifies a conserved MYB regulator of phenylpropanoid biosynthesis. Plant Cell 12:2383–2393 Butelli E, Licciardello C, Zhang Y, Liu J, Mackay S, Bailey P, Reforgiato-Recupero G, Martin C (2012) Retrotransposons control fruit-specific, cold-dependent accumulation of anthocyanins in blood oranges. Plant Cell 24(3):1242–1255 Chen WH (2017) The development of Phalaenopsis orchid industry in Taiwan. Orchid Digest 81-1:22–27 Chen WH, Hsu CY, Cheng HY, Chang H, Chen HH, Ger MJ (2011) Downregulation of putative UDP-glucose: flavonoid 3-O-glucosyltransferase gene alters flower coloring in Phalaenopsis. Plant Cell Rep 30:1007–1017 Chen WH, Kao YK, Tang CY (2014) Variation of the Genome Size among Phalaenospsis species using DAPI. J Taiwan Soc Hortic Sci 60:115–123 Chen WH, Kao YL, Tang CH (2013a) Method for producing polyploid plants of orchids. US Patent 8,383,881 Chen WH, Kao YK, Tang CY, Tsai CC, Lin TY (2013b) Estimating nuclear DNA content within 50 species of the genus Phalaenopsis Blume (Orchidaceae). Sci Hortic (Amsterdam) 161:70–75 Chiu LW, Zhou X, Burke S, Wu X, Prior RL, Li L (2010) The purple cauliflower arises from activa- tion of a MYB transcription factor. Plant Physiol 154(3):1470–1480 Christenson EA (2001) Phalaenopsis. In: Phalaenopsis: a monograph. Timber Press, Portland Chuang HT, Hsu ST, Shen TM (2008) Breeding barriers in yellow Phalaenopsis orchids. J Taiwan Soc Hortic Sci 54:59–66 Darwin C (1868) The variation of animals and plants under domestication, 1st edn. John Murray, London Feller A, Machemer K, Braun EL, Grotewold E (2011) Evolutionary and comparative analysis of MYB and bHLH plant transcription factors. Plant J 66:94–116 Griesbach RJ (1985) Polyploidy in Phalaenopsis orchid improvement. J Hered 76:74–75 Griesbach RJ (2005) A scientific approach to breeding blue orchid-exploring new frontiers in search of elusive flower colors. Orchids 74:376–379 Grotewold E (2006) The genetics and biochemistry of floral pigments. Annu Rev Plant Biol 57:761–780 Hichri I, Barrieu F, Bogs J, Kappel C, Delrot S, Lauvergeat V (2011) Recent advances in the tran- scriptional regulation of the flavonoid biosynthetic pathway. J Exp Bot 62:2465–2483 Hsiao YY, Jeng MF, Tsai WC, Chuang YC, Li CY, Wu TS, Kuoh CS, Chen WH, Chen HH (2008) A novel homodimeric geranyl diphosphate synthase from the orchid Phalaenopsis bellina lack- ing a DD(X)2-4D motif. Plant J 55:719–733 23 Phalaenopsis 625

Hsiao YY, Tsai WC, Kuoh CS, Huang TH, Wang HC, Wu TS, Leu YL, Chen WH, Chen HH (2006) Comparison of transcripts in Phalaenopsis bellina and Phalaenopsis equestris (Orchidaceae) flowers to deduce monoterpene biosynthesis pathway. BMC Plant Biol 6:14 Hsu CC, Chen WH (2015) The breeding achievements from Phalaenopsis equestris. Malayan Orchid Rev 49:41–47 Hsu CC, Chen YY, Tsai WC, Chen WH, Chen HH (2015a) Three R2R3-MYB transcription factors regulate distinct floral pigmentation patterning inPhalaenopsis spp. Plant Physiol 168:175–191 Hsu HF, Hsu WH, Lee YI, Mao WT, Yang JY, Li JY, Yang CH (2015b) Model for perianth forma- tion in orchids. Nature Plants 1:15046 Kao YY, Chiang SB, Lin TY, Hsieh CH, Chen YH, Chen WH, Chen CC (2001) Differential accu- mulation of heterochromatin as a cause for karyotype variation in Phalaenopsis orchids. Ann Bot 87:387–395 Koes R, Verweij W, Quattrocchio F (2005) Flavonoids: a colourful model for the regulation and evolution of biochemical pathways. Trends Plant Sci 10:236–242 Kuo PC, Wu TS (2011) Biosynthetic pathway of pigments in Phalaenopsis species. Orchid Biotechnol II:129–144 Lin S, Lee H, Chen W, Chen C, Kao Y, Fu Y, Chen Y, Lin T (2001) Nuclear DNA contents of Phalaenopsis sp. and Doritis pulcherrima. J Am Soc Hortic Sci 126:195–199 Ma H, Pooler M, Griesbach R (2009) Anthocyanin regulatory/structural gene expression in Phalaenopsis. J Am Soc Hortic Sci 134:88–96 OrchidWiz (2017) OrchidWiz Encyclopedia Version X3.3 May 2017 Database. OrchidWiz Database LLC, Ames Petroni K, Tonelli C (2011) Recent advances on the regulation of anthocyanin synthesis in repro- ductive organs. Plant Sci 181:219–229 Sasaki N, Nakayama T (2015) Achievements and perspectives in biochemistry concerning antho- cyanin modification for blue flower coloration. Plant Cell Physiol 56:28–40 Sattler MC, Carvalho CR, Clarinde WR (2016) The polyploidy and its role in plant breeding. Planta 243:281–296 Shindo S, Kamemoto H (1963) Karyotype analysis of some species of Phalaenopsis. Cytologia 28:390–398 Su V, Hsu BD (2003) Cloning and expression of a putative cytochrome P450 gene that influences the colour of Phalaenopsis flowers. Biotechnol Lett 25:1933–1939 Tanaka Y, Ohmiya A (2008) Seeing is believing: engineering anthocyanin and carotenoid biosyn- thetic pathways. Curr Opin Biotechnol 19:190–197 Tsai WC, Kuoh CS, Chuang MH, Chen WH, Chen HH (2004) Four DEF-like MADS box genes displayed distinct floral morphogenetic roles in Phalaenopsis orchid. Plant Cell Physiol 45:831–844 Wang J, Ming F, Han Y, Shen D (2006) Flavonoid-3′,5′-hydroxylase from Phalaenopsis: a novel member of cytochrome P450s, its cDNA cloning, endogenous expression and molecular mod- eling. Biotechnol Lett 28:327–334 Winkel-Shirley B (2001) Flavonoid biosynthesis. A colorful model for genetics, biochemistry, cell biology, and biotechnology. Plant Physiol 126:485–493 Yeh YC, Tsai YS, Shih CT, Huang P (2012) The breeding of scented flowers in Phalaenopsis. In: Orchid breeding in Taiwan. Published by Taiwan Orchid Breeders Society. pp 86–94 Zhang G, Lu S, Chen TA, Funk CR, Meyer WA (2003) Transformation of triploid bermuda grass (Cynodon dactylon × C. transvaalensis cv. TifEagle) by means of biolistic bombardment. Plant Cell Rep 21:860–864