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Cerium and Lanthanum Promote Floral Initiation and Reproductive Growth of Arabidopsis Thaliana

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Cerium and Lanthanum Promote Floral Initiation and Reproductive Growth of Arabidopsis Thaliana Plant Science 159 (2000) 117–124 www.elsevier.com/locate/plantsci Cerium and lanthanum promote floral initiation and reproductive growth of Arabidopsis thaliana Ya-Wen He, Chiang-Shiong Loh * Department of Biological Sciences, National Uni6ersity of Singapore, 14 Science Dri6e 4, Singapore 117543, Singapore Received 1 December 1999; received in revised form 2 June 2000; accepted 5 July 2000 Abstract The effects of cerium and lanthanum on the vegetative growth, floral initiation and reproductive growth of Arabidopsis thaliana were studied. Addition of cerium nitrate (0.5–10 mM) or lanthanum nitrate (0.5–50 mM) to the culture medium significantly increased the lengths of primary roots, but had no significant effects on the number of rosette leaves produced per plant, plant heights and dry weights during the vegetative growth stage (17 days after seed germination). The percentage of plants bolted was significantly increased with the addition of 0.5–10.0 mM cerium nitrate or lanthanum nitrate. The combination of 0.5 mM cerium nitrate and 0.5 mM lanthanum nitrate was found to be most effective on the induction of floral initiation. The height, dry weight and average number of flower numbers of 35-day-old plants growing in media containing cerium nitrate or/and lanthanum nitrate (0.5–10.0 mM) were found to be significantly higher than those in the control medium. The endogenous levels of cytokinins (zeatin riboside, dihydrozeatin riboside and isopentenyl adenosine) and carbohydrates (sucrose, glucose and fructose) in leaf and root tissues of plants growing in the medium supplemented with 0.5 mM cerium nitrate and 0.5 mM lanthanum nitrate were not significantly different from those of plants in the control medium. Application of 0.5 mM cerium nitrate and 0.5 mM lanthanum nitrate enhanced the effects of 10−6 M IPA on root growth, plant height and flowering. The role of cerium and lanthanum in promoting floral initiation and reproductive growth and the possibility of developing non-hormonal flowering promoting agents are discussed. © 2000 Elsevier Science Ireland Ltd. All rights reserved. Keywords: Arabidopsis thaliana; Cerium; Lanthanum; Cytokinin; Flowering 1. Introduction REEs makes it imperative that more detailed in- formation on the effects of REEs on biological The rare earth elements (REEs) comprise a systems be available. group of 15 trivalent metallic elements with similar The effects of REEs, especially lanthanum, in chemical properties. They normally occur as basic different animal tissues have been studied exten- oxides and phosphate complexes in nature [1,2]. sively [1]. Most of the work carried out on REEs Since the introduction of ion exchange techniques, in animal systems was based on the use of REE the separation of the rare earth elements from ion as a substitute or antagonist for Ca2+ to their ores and from one another has become prac- monitor the movement of calcium and water, and tical and many new uses of REEs have been to investigate the role of calcium in muscle and developed [3]. The widespread industrial usage of nerve activity [4]. Abbre6iations: BA, benzyladenine; Ce, Cerium; DHZR, dihy- The results from limited studies on the effect of drozeatin riboside; IPA, isopentenyl adenosine; La, Lanthanum; MS REEs on plant growth are conflicting. Early re- medium Murashige and Skoog (1962) medium; Nd, Neodymium; ports indicated that the REE were inhibitory to REEs, rare earth elements; ZR, zeatin riboside. 3+ 3+ * Corresponding author. Tel.: +65-874-2916; fax: +65-779-5671. plant growth. For example, La and Nd were E-mail address: [email protected] (C.-S. Loh). found to inhibit elongation of oat coleoptile sec- 0168-9452/00/$ - see front matter © 2000 Elsevier Science Ireland Ltd. All rights reserved. PII: S0168-9452(00)00338-1 118 Y.-W. He, C.-S. Loh / Plant Science 159 (2000) 117–124 tions [5]. Colloidal lanthanum caused an almost drate (La(NO3)3·6H2O, Sigma) were dissolved in complete inhibition of cell division and root elon- Mili-Q water and sterilized by membrane filtra- gation in the root tips of barley plants [6]. La3+ tion (Millipore, 0.45 mm) and stored at room had been shown to inhibit root elongation of temperature in the dark. Stock solutions of wheat [7,8]. More recent reports, however, cerium nitrate and lanthanum nitrate were added demonstrated some positive effects of REEs on to the autoclaved basal medium prior to dispens- plant growth. Diatloff et al. [9] reported that ing into Magenta GA7® containers (Magenta corn root growth increased significantly with the Corp., USA). Seeds were germinated in the dark applications of cerium (0.63 mM) and lanthanum and 2-day-old seedlings were placed under 16 h (0.63 mM). Applications of lanthanum and photoperiod (54 mmol−1m−2s−1 provided by cerium were also reported to increase spike pro- Cool White fluorescent lamps) at 2592°C. duction in wheat [10]. In pot trials, applications of cerium sulphate (up to 100 mg/kg) enhanced 2.2. Growth measurements root and shoot growth of Phaseolus radiatus and Brassica pekinensis [11]. Lengths of primary roots were scored 10 days Results from field trials were also inconsistent. after seed germination. The number of leaves The increase in crop yield reported by workers produced per plant was scored 17 days after seed from China ranged between 8 and 50%, with the germination. Plant heights and dry weights were common response being of the order of 8–15% scored 17 and 35 days after seed germination, [12,13]. However, no response was found by respectively. Dry weights were taken by drying spraying and seed dressing of a summer fodder 100 plants in an oven (55°C) for 1 week. Floral crop (Brassica sp.) with REEs in a field trial initiation was recorded when the plant bolted carried out in Australia (cited in [4]). In view of with at least 1 cm long inflorescence stalk. this, it is essential to study and elucidate the effects of REE on essential stages of growth and 2.3. Extraction and determination of endogenous development of a model plant species such as cytokinins and carbohydrates Arabidopsis thaliana. In this report, we investi- gated the effects of cerium (Ce) and lanthanum Approximately 1 g fresh weight of tissues was (La) on vegetative and reproductive growth of A. homogenized in 4 ml of 80% ethanol followed by thaliana and correlated some of the responses to 1 h incubation at 4°C. After centrifugation at increased sensitivity of cell to plant growth regu- 1670×g for 3 min, the supernatant was trans- lators. ferred to another centrifuge tube. The tissues were re-extracted with 2 ml of 80% ethanol, and the supernatant was pooled together after cen- trifugation. The extracts were vacuum evaporated 2. Materials and methods at 4°C (Eppendorf Concentrator 5301) to 1/4 volume and then stored at −20°C after filter- 2.1. Plant materials, culture media and growth sterilization (Milipore, 0.2 mm). The analysis of conditions zeatin riboside (ZR), dihydrozeatin riboside (DHZR), and isopentenyl adenosine (IPA) were Seeds of A. thaliana L. Heynh cv. Columbia performed by immunoassay detection kits (Sigma (LEHLE SEEDS, USA) were surface sterilized Chemical Company) according to the protocols by soaking in 75% alcohol for 30 s and followed provided by the manufacturer. All hormonal lev- by 15% Clorox® for 15 min. The seeds were then els were expressed in terms of pmol per gram rinsed five times in sterilized water prior to cul- fresh weight (pmol/g.f.wt). The analysis of su- ture. The 1/4 strength Murashige and Skoog crose, glucose and fructose were performed by medium [14] was used for seed germination and assay kits (Sigma Chemical Company) according as basal medium. The pH of the medium was to the protocols provided by the manufacturer. adjusted to 5.8 before agar (Difco, 0.8%) was The levels of sucrose were expressed in term of added. All media were autoclaved for 20 min at mg per gram fresh weight (mg/g.f.wt). The levels 121°C. Cerium nitrate hexahydrate (Ce(NO3)3 of glucose and fructose were expressed in term of m m ·6H2O, Sigma) and lanthanum nitrate hexahy- g per g fresh weight ( g/g.f.wt). Y.-W. He, C.-S. Loh / Plant Science 159 (2000) 117–124 119 3. Results tions of cerium nitrate or lanthanum nitrate (0.5– 2.5 mM) significantly increased the percentages of 3.1. Effects on 6egetati6e growth plants bolted on days 20 and 23. For example, 69.3% of the plants growing in the medium with The vegetative and reproductive growth of A. 2.5 mM cerium nitrate bolted on day 23 whereas thaliana plant was separated. Prior to bolting, an only 39.4% bolted in basal medium. About 65.7% A. thaliana plant consisted of a rosette of small of the plants in the medium containing 0.5 mM leaves with a main hypocotyl. In the present lanthanum nitrate bolted on day 23 (Table 2). study, results showed that additions of 0.5–50 mM The percentages of plants bolted on day 27 were cerium or lanthanum had no significant effects on not significantly different between media with or the vegetative growth in terms of height and dry without cerium nitrate or lanthanum nitrate weight. Cerium and lanthanum also had no sig- (Table 2). nificant effect on the average number of rosette None of the plants growing in basal medium leaves 17 days after seed germination (data not bolted on day 19. Additions of certain combina- shown). Additions of higher concentrations of tions of cerium nitrate and lanthanum nitrate (e.g. cerium or lanthanum inhibited the vegetative 0.5+0.5 mM, 0.5+1.0 mM, 1.0+0.5 mM, 1.0+ growth (data not shown).
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