Stems of Allium cepa L. contain starch

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1 New Phytol. (1994), 128, Stems of Allium cepa L. contain starch BY M. ERNST AND G. BUFLER Institut fiir Obst-, Gemiise- und Weinbait, Universitdt Hohenheim, D Stuttgart, Germany {Received 20 April 1994; accepted 6 July 1994) SUMMARY Stems oi Allium cepa L. were prepared for histochemical analysis and stained by periodic acid-schiff reaction and Lugol's iodine for detecting starch grains (amyloplasts). Starch was found in the root cap of the root initials, around the central cylinder vascular tissues of the stem and in the primary thickening meristem during sprouting hut not during dormancy. Key words: Allium cepa (onion), starch, dormancy. INTRODUCTION The non-structural carbohydrates in onion {Allium cepa L.) include glucose, fructose, sucrose and a series of oligosaccharides, the fructans. Numerous investigations have been conducted on fructans in onion, including their distribution (Darbyshire & Henry, 1978) and changes during storage (Darbyshire, 1978; Rutherford & Whittle, 1982; Suzuki & Cutcliffe, 1989). There are few reports of starch in onion, but it has been reported that starch is usually not detectable in the vegetative tissues of this crop (Gates & Simpson, 1968; Darbyshire & Steer, 1990). However, starch grains were reported in the gravitational base of columella cells in onion roots (Moore, 1985). Wilson, Oross & Lucas (1985) reported that chloroplasts in the leaf bundle-sheath cells of 10-wk-oId onion seedlings contain starch and discussed their function in relation to carbohydrate pools and translocation. Komochi (1990) reviewed sprouting, respiration rates, rooting, plant hormones and carbohydrates with the objective of finding an indicator of onion bulb dormancy (the period during which visible sprout growth is not detectable), but none of these parameters was useful for this purpose. Histological investigations on Helianthus tuberosus L. showed that starch is detectable in the tuber during tuber development, fructan synthesis and the beginning of sprouting, but not during tuber dormancy (Ernst, 1991). This pattern of the occurrence of starch may also be a feature of onion bulbs, and if so, the absence of starch may indicate dormancy. To test this hypothesis, onion stems wei-e analyzed during bulb development and storage and investigated for the presence of starch in two consecutive years (1992, 1993). Only data from 1993 are presented. MATERIALS AND METHODS Plant materials Onions cv. Super Bear were field-seeded on 14 April 1993 in Stuttgart-Hohenheim. On 8 August 1993, when 80 % of the onion leaves had fallen, bulbs were harvested, cured and stored in open boxes at C and 40-60% r.h. Histochemical analysis The basal plate of onion rather than the bulb forming leaf bases was used in this study because it contains meristematic tissue and is located where rooting and sprouting commence after dormancy. At 7 d intervals, starting on 26 May 1993 and continuing until 26 January 1994, basal plates were fixed in EPA 70 (5% (v/v) of formaldehyde solution 37% GR, 5 % (v/v) of propionic acid, 90 % (v/v) of ethanol 70% GR). Eollowing dehydration in an ethanol series, and embedding in glycolmethacrylate-methyl methacrylate (Ruddell, 1967) microscopic sections (5 /im) from collected samples were stained with periodic acid-schiff reaction and Lugol's iodine (Chayen & Bitensky, 1991) and tested for starch. Determination of dormancy and sprout growth {'leaf ratio') At each date of sampling 10 bulbs were dissected into scale and sprout leafs. The youngest sprout leaf (leaf initial) which was still macroscopically discernible was isolated and its 'leaf ratio' determined. According to Heath & Hollies (1965) the 'leaf ratio' is defined as the ratio of leaf blade length to leaf sheath length; the leaf pore marks the junction between leaf blade and sheath. After dormancy when

2 404 M. Ernst and G. Bufler Ri St Figure 1. Starch grains (St) in the root cap initials (RI) within the stem of Alliuni cepa ( x 250). st Figure 2. Starch grains (St) in cells around the vascular tissue (VT) of the stem of AUium cepa ( x 400). the shoot resumes active growth and the leaf initials elongate, the leaf ratio increases. RESULTS AND DISCUSSION Our studies revealed the occurrence of starch in three different tissues of the onion stem, in the root tip initials, around the vascular tissue, and in the primary thickening meristem. In addition to findings of starch grains by Moore (1985) in the root tip caps outside the basal plate, our studies show that starch is also present in the root tip initials inside the basal plate at the time of rooting (Fig. 1). The central cylinder is congested with vascular bundles and leaf traces. Leaf traces are collateral and stem bundles are amphivasal in vascular tissue arrangement (DeMason, 1990). In cells around these

3 405 Starch in Allium cepa Figure 3. Starch grains forming a band in the primary thickening meristem of the stem of Allium cepa ( x 160). Figure 4. Starch grains in the primary thickening meristem of the stem of Allium cepa by SEM ( x 18000). vascular tissues, starch grains were detected in a few samples collected at different times (Fig. 2). The primary thickening meristem of the stem can be distinguished as a cambium-like zone consisting of tangentially flattened cells located between the cortex and the central cylinder (DeMason, 1979). It is the site of cell division activity during the primary stem thickening process in onion (DeMason, 1980). Bands of starch grains (Fig. 3), interrupted by root initials, were detected in the primary thickening nieristem of samples collected from 26 May 1993 until 21 July 1993 (19 d before harvest), and after 13 October 1993 (66 d after harvest) until 26 January 1994 (end of storage). Results from 1992 showed a similar pattern. The diameter of the globular amyloplasts was less than 2 //.m (Fig. 4). During the period of bulb dormancy (middle of July until end of October, 1993), as determined by recording of the ' leaf ratio', starch was not (except in trace amounts), detectable in the primary thickening meristem (Table 1). The absence of starch in the primary thickening meristem therefore may be indicative of onion bulb dormancy. We are currently studying the effect of dormancy period length on the pattern of starch occurrence in the primary thickening meristem as well as whether or not starch in

4 406 M. Ernst and G. Bufler Table 1. Occurrence of starch in cells of the primary thickening meristem of onion stem during bulb development and dormancy as determined by 'leaf ratio' Date of sampling (1993) 30 Jun. 21 Jul. 18 Aug. 13 Oct. 10 Nov. 24 Nov. Bulb diam. (mm) ' Leaf ratio' 0-9 0' O-5 2-H ±l-2 Occurrence of starch ' Sprout growth resumed, i.e. end of dormancy,, traces of starch;, starch present. cells of the primary thickening meristem plays a role in osmoregulation during periods of high metabohc activity. ACKNOWLEDGEMENTS Thanks are due to Mrs Hepp (Institut fur Physik, Universitat Hohenheim) for her expert assistance with the electron microscopy. Many thanks to Boyd G. Penney for critical i-evision of the English text. REFERENCES ChayenJ,Bitensky L Practical histochemistry. Chichester: John Wiley & Sons Ltd. Darbyshire B Changes in the carbohydrate content of onion bulbs stored for various times at different temperatures. Journal of Horticultural Science 53: Darbyshire B, Henry RJ The distribution of fructans in onions. New Phytologist 81: Darbyshire B, Steer BT Carbohydrate biochemistry. In: Brewster JL, Rabinowitch HD, eds. Onions and allied crops, vol. III. Boca Raton: CRC Press, DeMason DA Histochemistry of the primary thickening meristem in the vegetative stem of Allium cepa L. American Journal of Botany 66: DeMason DA Localisation of cell division activity in the primary thickening meristem in Allium cepa L. American Journal of Botany 67: DeMason DA Morphology and anatomy of Allium. In: Rabinowitch HD, Brewster JL, eds. Onions and allied crops, vol. I. Boca Raton: CRC Press, Ernst M Histochemische Untersuchungen auf Inulin, Starke und Kallose bei Helianthus tuberosus L. (Topinambur). Angewandte Botanik 65: Gates JW, Simpson GM Tbe presence of starch and alpha-amylase in tbe leaves of plants. Canadian Journal of Botany 46: Heath OVS, Hollies MA Studies in tbe physiology of tbe onion plant. VI. A sensitive morphological test for bulbing and its use for detecting bulb development in sterile culture. Journal of Experimental Botany 16: Komochi S Bulb dormancy and storage pbysiology. In: Rabinowitcb HD, Brewster JL, eds. Onions and allied crops, vol. I. Boca Raton: CRC Press, Moore R Calcium movement, graviresponsiveness and the structure of columella cells and columella tissues in roots of Allium cepa L. Annals of Botany 56: Ruddell CL Hydroxyethyl methacrylate combined with polyethylene glycol 400 and water, an embedding medium for 1-2 micron sectioning. Stain Technology 42: Rutherford PP, Whittle R The carbohydrate composition of onions during long term cold storage. Joi/r/m/ of Hortieuttural Science 57: Suzuki M, Cutcliffe JA Fructans in onion bulbs in relation to storage life. Canadian Journal of Plant Science 69: Wilson C, Oross JW, Lucas WJ Sugar uptake into Allium eepa leaf tissue: an integrated approach. Planta 164:

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