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Differences in Floral Development Between Lycoris Radiata And R ESEARCH ARTICLE doi: 10.2306/scienceasia1513-1874.2020.032 Differences in floral development between Lycoris radiata and Lycoris sprengeri a,† b,† c d a a,e, Junhuo Cai , Junjun Fan , Xuying Wei , Donglin Zhang , Jiajia Ren , Lu Zhang ∗ a College of Landscape and Art, Jiangxi Agricultural University, Nanchang 330045 China b College of Horticulture, Jinling Institute of Technology, Nanjing 210037 China c College of Art, Jiangxi Finance and Economics University, Nanchang 330032 China d Department of Horticulture, University of Georgia, Athens, GA 30602 USA e Collaboration Innovation Center of Jiangxi Typical Trees Cultivation and Utilization, Jiangxi Agricultural University, Nanchang 330045 China ∗Corresponding author, e-mail: [email protected], Cai J. and Fan J. contributed equally to this paper. Received 12 Sep 2019 Accepted 21 Mar 2020 ABSTRACT: Lycoris radiata and Lycoris sprengeri are two typical plants of the genus Lycoris. They have different leafing and flowering patterns. To understand the habits of Lycoris species and regulate the flowering period effectively, we compared flower bud development of these two species using phenological, anatomical, and physiological measurements. Our results showed there were significant differences in phenology between the two species. The sprouting stage of L. radiata took place in autumn, the leaf growth period lasted seven months, dormancy lasted approximately 100 days before flowering, and there was no dormancy period after flowering. In comparison, the sprouting period of L. sprengeri was delayed until the spring of the following year, the leaf growth stage lasted approximately 3 months, the flowering stage took place from Jul–Aug, and dormancy lasted 1–2 months before flowering and for another 6 months after flowering. The flower bud differentiation processes of the two species were similar. During the entire flower bud differentiation process, the soluble sugar and protein contents, as well as the superoxide dismutase activity, in L. sprengeri bulbs were significantly higher than in L. radiata bulbs, whereas peroxidase activity was significantly lower. KEYWORDS: phenology, flower bud differentiation, flower development INTRODUCTION years, the study of Lycoris has promoted its produc- tion and use in gardening, and 368 hm2 of L. radiata There are 20 species of genus Lycoris worldwide. have been artificially cultivated in Jiangxi [5]. Lycoris plants are unique herbs in Southeast Asia, The life cycle of Lycoris plants can be di- and are mainly produced in China (15 species, vided into aboveground and underground stages. 2 varieties) and Japan; few are grown in Burma and The aboveground stages include flowering and leaf North Korea 1, 2 . Around 10 species are widely [ ] sprouting [2], whereas the underground stages are used in gardening and medicine [3]. There are 12 dominated by bulb growth and dormancy. Lycoris species that are unique to China [4], which are com- plants are divided into two types based on autumn monly found south of the Yangtze River and consist sprouting (mid-end Oct) or spring sprouting (early primarily of temperate species [3]. Lycoris plants Feb). In general, L. radiata (L’Hér.) Herb. is the type have diverse flower colors and bloom in midsummer of sprouting, which blooms in early autumn [6]. or early autumn. In addition, these plants are toler- L. sprengeri Comes Ex Baker is thought to be the ant to drought and nutrient deficiency and are also type of spring sprouting, which blooms in summer. less sensitive to pests and diseases. Therefore, these However, other studies have found that the leaves plants are superior materials for bulb flowers and of L. sprengeri continue to emerge from autumn to ground cover in gardens. Breeding work of Lycoris spring (the percentage of leave-sprouting seedlings plants started in the 1940s in various countries, but in autumn was less than 50%, and then they con- the first hybrid L. elegans (L. sprengeri L. chinensis) tinued to emerge in the following spring). Leaf bud was obtained in China in the 1970s [×1]. In recent www.scienceasia.org 2 ScienceAsia 46 (2020) differentiation in Lycoris plants begins around the was 15 cm 20 cm. Five sampling points in every end of Mar and is completed in Sep. The flower observation× point were used to select 5 plants for bud differentiation processes of the two types of observation and recording of phenological traits Lycoris plants are generally similar. Flower bud dif- during the entire growth period [7]. Photographs ferentiation occurs in underground bulbs in Lycoris of the plants were obtained (from leaf sprouting to plants, during which almost no leaves are present. flower withering). The starting time of flower bud differentiation in The phenological record cycle was from 1 Sep Lycoris plants is easily affected by environmental 2013 to 20 Oct 2014. The plants were observed factors such as temperature. For this reason, various once every 5 days. The number of sprouted leaves, researchers have identified different distinguishing leaf length and width, leaf tip yellowing, leaf wither- features for the flower bud differentiation stage in ing, leaf withered, scape emergence, scaping, flow- the same plants [6]. ering, and flower withering were recorded. The Despite the increase in the number of studies on division criteria of different phenological periods the regulation of flowering in Lycoris plants, system- were as follows. Leaf sprouting period was from atic investigations on flower development are still 5% to 85% of the plants with sprouting leaves. limited. The present study conducted a comparative Greening period was from 85% of the plants with analysis of the phenological characteristics, mor- sprouting leaves to 5% of the plants with yellow phological anatomy-related characteristics of flower leaves. Defoliation period (leaf yellowing and with- bud differentiation, and physiological characteris- ering) was from 5% of the plants with yellow leaves tics of flower development for two species of dif- to 85% of the plants without leaves. Dormancy ferent sprouting types: L. radiata and L. sprengeri. period (leafless) was from 85% of the plants without The results of this study may serve as a foundation leaves to 5% of the plants with sprouting leaves or for future studies on the hybridization and breeding scape sprouting. Flowering period (scape sprouting of Lycoris plants, as well as provide insight for and blooming) was from scape sprouting to flower understanding the diversity of growth rhythms and withering. for investigations on the regulation of flowering in Lycoris plants. Paraffin sections of bulb shoots MATERIALS AND METHODS From 10 Mar–25 Jul 2014, L. radiata and L. spren- Overview of study materials and site geri bulbs were collected once every 5 days for a total of five times. We randomly sampled 1 bulb at L. radiata (L’Hér.) Herb. and L. sprengeri Comes each observation point. The bulbs were collected Ex Baker were obtained from the experimental regularly, and the central flower bud was removed flower nursery in Jiangxi Agricultural University and washed with deionized water before fixing in a (Nanchang, China, 28.76°N, 115.83°E). L. radiata mixed fixative solution (90 ml of 70% alcohol 5 ml generates quadrennial bulbs, which are obtained + glacial acetic acid 5 ml of formalin, FAA) for more by asexual reproduction and have a diameter of + than 24 h. Routine paraffin sections (8 m thick) 3.5 0.1 cm. The soil used was red soil that was de- µ were prepared for flower bud morphological obser- veloped± from Quaternary red clay parent material. vation 7 , with slight modifications. The sections In the 0–40 cm soil layer, the particle distribution [ ] were stained with iron (II) sulfate-hematoxylin and was as follows: 12.12, 51.63, 35.87, and 0.38% for examined and photographed using a stereoscopic soil depths of <0.002, 0.002–0.05, 0.05–2.00, and microscope (DM2500, Leica, Germany). >2.00 mm, respectively. Soil organic matter content was 13.5 g/kg. Soil nitrogen, phosphorus, and potassium contents were 0.84, 0.24, and 24.9 g/kg Physiological measurements of bulbs respectively, and the pH was 6.43. Nine Lycoris (triplicates, 3 plants per replicate) Experimental methods plants were randomly selected every 5 days and their bulbs were extracted. After washing with tap Phenological observations water and drying, the first 2–4 layers were removed, Fixed plant observations were performed; 5 obser- crushed, pooled, and used for measuring various vation points for each species were used. The area physiological markers. Soluble sugar content was of each observation point was 300 300 cm2, and measured by anthrone colorimetry. Soluble protein 300 Lycoris plants were planted. The× row spacing content was determined using Coomassie blue (G- www.scienceasia.org ScienceAsia 46 (2020) 3 250) staining, and nitroblue tetrazolium (NBT) pho- year), warm springs (11.8–23 °C), burning hot sum- toreduction was employed to measure superoxide mers (26–29.6 °C) and cold winters (2.8–8.1 °C), dismutase (SOD) activity (1 enzyme unit (U) is long summers and winters, and short springs and equivalent to a 50% inhibition of NBT photore- autumns (13.9–25.3 °C). In Nanchang, Aug–Sep is duction). Peroxidase (POD) activity was measured the flowering period and leaves start to sprout in Oct using guaiacol colorimetry. after the flowers have withered. The plants enter a dormant period in May of the following year after Data analysis the leaves have defoliated. The mean and standard deviation of each marker The leaves of L. sprengeri are banded and are was analyzed using Excel 2013. One-way ANOVA 30 cm in length, 1.0 cm in width, green in color, (Duncan’s test) in SPSS 17.0 was used for the and have blunt tips. Scape height is approximately comparison of the same factor at different levels of 60 cm. There are 2 bracts that are 3.5 cm in length significance (p < 0.01).
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