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Development, Progress and Future Prospects in Cryobiotechnology of Lilium Spp

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Development, Progress and Future Prospects in Cryobiotechnology of Lilium Spp Li et al. Plant Methods (2019) 15:125 https://doi.org/10.1186/s13007-019-0506-9 Plant Methods REVIEW Open Access Development, progress and future prospects in cryobiotechnology of Lilium spp. Jing‑Wei Li1,2, Xiao‑Chen Zhang1, Min‑Rui Wang1, Wen‑Lu Bi1,3, M. Faisal4, Jaime A. Teixeira da Silva5, Gayle M. Volk6 and Qiao‑Chun Wang1* Abstract Lilium is one of the most popular fower crops worldwide, and some species are also used as vegetables and medi‑ cines. The availability of and easy access to diverse Lilium genetic resources are essential for plant genetic improve‑ ments. Cryopreservation is currently considered as an ideal means for the long‑term preservation of plant germplasm. Over the last two decades, great eforts have been exerted in studies of Lilium cryopreservation and progress has been made in the successful cryopreservation of pollen, seeds and shoot tips in Lilium. Genes that exist in Lilium, including those that regulate fower shape, color and size, and that are resistant to cold stress and diseases caused by fungi and viruses, provide a rich source of valuable genetic resources for breeding programs to create novel cultivars required by the global foriculture and ornamental markets. Successful cryopreservation of Lilium spp. is a way to preserve these valuable genes. The present study provides updated and comprehensive information about the devel‑ opment of techniques that have advanced Lilium cryopreservation. Further ideas are proposed to better direct future studies on Lilium cryobiotechnology. Keywords: Cryopreservation, Genetic stability, Lily, Pollen, Seeds, Shoot tips Highlights America, and 10 Lilium species to European and Cauca- sus [1]. • Availability of diverse genetic resources is essential Lilium is globally grown as one of the most important for Lilium genetic improvements; cut and pot fower and garden crops, mainly due to large, • Cryopreservation is an ideal means for the long-term fragrant, multi-colored fowers with a long shelf-life [3, preservation of plant germplasm; 4]. Lilium species and hybrids are also used as garden • Tis review provides recent advances in the said feld; plants [3]. Te bulbs of some Lilium species including • Future prospects are proposed to direct further stud- L. davidii var. unicolor, L. brownii var. viridulum and L. ies. lancifolium are widely consumed as vegetables [5] due to their high levels of protein, amino acids, minerals and dietary fber [6, 7]. Other Lilium species such as L. regale, Background L. concolor, L. pumilum, L. davidii var. unicolor and L. Lilium, a perennial bulb plant, includes approximately lancifolium are traditional Chinese medicines [8], and 100 species [1]. About 55 species and 32 varieties of Lil- they contain biochemical compounds such as alkaloids, ium are native to China [2], 21 Lilium species to North polysaccharides, saponin and colchicine that have anti- oxidant activities [6, 9]. Lilies have been grown as vegeta- bles and medicine for at least 2000 years in China [10]. A wide diversity of Lilium genetic resources has pro- *Correspondence: [email protected] 1 State Key Laboratory of Crop Stress Biology in Arid Region, College vided valuable genes for breeding of novel cultivars. of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, Tere are more than 300 new registrations of Lilium cul- People’s Republic of China tivars each year and approximately 9500 cultivars in total Full list of author information is available at the end of the article © The Author(s) 2019. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creat iveco mmons .org/licen ses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Li et al. Plant Methods (2019) 15:125 Page 2 of 12 globally [11]. Tere is a demand for new, novel lily culti- Shoot tips vars from the production industry [4, 12, 13]. A shoot tip refers to an apical dome and a few leaf pri- Te availability of and easy access to diverse plant mordia that are composed of undiferentiated cells genetic resources are essential to provide the valuable capable of continuous cellular division and giving rise genes necessary to develop novel cultivars. Extensive to a shoot [39]. For vegetatively propagated plants for explorations and collections, heavy grazing, forest plant- which specifc allelic combinations must be preserved, ing and urban expansions have threatened wild Lilium shoot tips are preferred over cell suspensions, seeds and species [14, 15]. Global climate warming, which has embryogenic tissues for preservation of plant germplasm, increased the mean temperature of the Earth’s surface by because specifc genotypes are conserved [20, 21]. Lil- approximately 0.8 °C from the early 1900s to 2011 [16], ies are vegetatively propagated in the foriculture indus- has further worsened the situation [15, 17]. L. tsingtau- try because specifc cultivars must be maintained and ense, L. polyphyllum, L. pomponium and L. maculatum therefore, shoot tips are often the preservation target var. bukosanense are listed as endangered species in for cryopreserving Lilium germplasm. A list of success- China [14], Italy [18], India [15] and Japan [19] respec- ful cryopreservation of Lilium shoot tips is presented in tively. As a result, it is necessary to preserve and protect Table 1. Lilium genetic resources. Cryopreservation is a method whereby plant genetic resources can be conserved in liquid nitrogen (LN, Droplet‑vitrifcation − 196 °C) or liquid nitrogen vapor (LNV, approx. − 165 Vitrifcation refers to a physical process by which highly to − 190 °C) [20–22]. Once stored in liquid nitrogen con- concentrated and viscous cryoprotectant mixtures form ditions, cellular divisions of the samples cease, and theo- metastable glasses after rapidly cooling to LN, without retically, plant materials can be maintained for extended the formation of ice crystals between and inside plant lengths of time [20–23]. Once placed into long-term cells [40]. Vitrifcation cryopreservation is among the storage, less space and fewer resources are required to most often used methods for cryopreserving plant germ- maintain secure collections, compared to feld or in vitro plasm [21, 22, 41]. Droplet-vitrifcation combines the collections [20–22]. Cryopreservation eliminates the risk advantages of droplet freezing with vitrifcation, thus of losing collections as a result of pathogen attacks and allowing samples to obtain rapid cooling and warming unexpected environmental conditions in feld collections rates, increasing the survival of cryopreserved samples and reduces the risk of losing cultures by contamina- [41, 42]. Droplet-vitrifcation is considered to be a robust tion and of genetic variations by repeated subcultures in means for the cryopreservation of Lilium shoot tips [32, in vitro collections [20–22]. 34–36, 38, 43]. Tis review provides updated and comprehensive Chen et al. [36] described a droplet-vitrifcation for Lil- information on the developments and advances in Lilium ium shoot tips. In their study, 2-months old in vitro lily cryopreservation. In additional to preserving genetic stock shoots were cold-hardened at 4 °C for 1 week under resources, successful cryopreservation provides a method a 16-h photoperiod of 35 µmol m−2 s−1 light intensity. to conserve valuable genes in Lilium spp. that regulate Shoot tips excised from the cold-hardened shoots were fower shape, color and size, and that are resistant to cold precultured in liquid Murashige and Skoog [44] medium stress and diseases caused by fungi and viruses. Pros- (MS) supplemented with 0.3 M sucrose for 2 days. Pre- pects are proposed to direct further studies on Lilium cultured shoot tips were loaded with a loading solution cryobiotechnology. composed of MS supplemented with 2.0 M glycerol and 0. 4 M sucrose for 20–40 min at 22 °C, and exposed to plant vitrifcation solution 2 (PVS2) [45] for 90–120 min Cryopreservation at 0 °C. PVS2 contains 30% (w/v) glycerol, 15% (w/v) eth- Bouman and De Klerk [24] were the frst to cryopre- ylene glycol, 15% (w/v) dimethylsulfoxide (DMSO) and serve shoot tips of L. speciosum with about 8% survival. 0.4 M sucrose in MS (pH 5.8) [45]. Dehydrated shoot tips Since then, various cryopreservation methods have been were transferred to 1.5-μL PVS2 droplets placed on ster- described for Lilium pollen [25], seeds [26–29] and shoot ile aluminum foil strips (5 × 30 mm), each strip carrying tips [4]. Te major cryoprotocols available for Lilium 5–6 shoot tips. Te aluminum foil strips were folded to include dehydration [26, 28, 29], encapsulation–dehydra- enclose the shoot tips and directly immersed into LN. For tion [29, 30], encapsulation–vitrifcation [30], vitrifca- thawing, frozen aluminum foil strips were removed from tion [30–33], and droplet-vitrifcation [34–38]. LN and immediately placed in 1.2 M sucrose at room Major steps of Lilium cryopreservation for shoot tips, temperature for 15 min, followed by post-culture on a pollen and seeds are provided in Fig. 1. recovery medium for shoot regrowth. Recovery medium Li et al. Plant Methods (2019) 15:125 Page 3 of 12 1 Shoot tips Preculture Loading Dehydration by PVS2 PVS2 droplet A Dehydration by 30% In vitro 2 PVS3 droplet stock plants PVS3, followed by 100% PVS3 2, 3, 4, 5 1, 2, 3, 4, 5 3 Dehydration Cryostorage Cold-hardening Shoot tips Preculture Loading Warming by PVS2 in LN 4 Dehydration Encapsulation by PVS2 1, 2, 3, 4 5 4, 5 Dehydration Post-culture Unloading 5 by air drying for recovery B Dry anthers on Extract anthers Package pollen in Cryostorage Germination Flowers paper at 25±2°C Warming from flowers aluminum foil in LN or TTC test until pollen release 1 Encapsulation Dehydration by air-drying C 1, 2, 3 2 Cryostorage Freshly collected seeds Cold storage Incubation in 30% glycerol Warming Germination in LN Dehydration over silica gel 3 Fig.
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