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Allelopathic effect of ginger on seed


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Scientia Horticulturae 116 (2008) 330–336 www.elsevier.com/locate/scihorti

Allelopathic effect of ginger on seed germination and seedling growth of soybean and chive
Chun-Mei Han, Kai-Wen Pan *, Ning Wu, Jin-Chuang Wang, Wei Li
Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China Received 28 March 2007; received in revised form 28 December 2007; accepted 10 January 2008

Abstract The rhizome, stem and leaf aqueous extracts of ginger were assayed at 10, 20, 40, and 80 g l?1 for their effects on seed germination and early seedling growth of soybean and chive. All aqueous extracts at all concentrations inhibited seed germination, seedling growth, water uptake and lipase activity of soybean and chive compared with the control, and the degree of inhibition increased with the incremental extracts concentration. The degree of toxicity of different ginger plant parts can be classi?ed in order of decreasing inhibition as stem > leaf > rhizome. The results of this study suggest that rhizome, stem and leaf of ginger contain water-soluble allelochemicals which could inhibit seed germination and seedling growth of soybean and chive. The rhizome is the main harvested part of ginger. The residue (mainly stems and leaves) of the ginger plant should be removed from the ?eld so as to diminish its inhibitory effect. Further work is needed to specify and verify the allelochemicals produced by this plant. The results of this study suggest that ginger allelochemicals are heterotoxic, and thus intercropping should not be practiced using ginger. # 2008 Elsevier B.V. All rights reserved.
Keywords: Ginger; Zingiber of?cinale Rosc.; Soybean; Glycine max (L.) Merr.; Chive; Allium schoenoprasum L.; Germination; Seedling growth; Water uptake; Lipase activity; Allelochemicals

1. Introduction Ginger (Zingiber of?cinale Rosc.) is an important horticultural crop in tropical Southeast Asia. It produces a pungent, aromatic rhizome that is valuable all over the world either as a spice or herbal medicine (Guo and Zhang, 2005). Many scholars have reported that ginger has the function of relieving the severity of nausea and vomiting during pregnancy (Vutyavanich et al., 2001; Portnoi et al., 2003). Additionally, it has been reported that crude extract of ginger rhizome can reduce rat paw and skin edema (Penna et al., 2003). However, ginger yields are low when this species is cultivated consecutively for years on the same land and rotated with other crops for at least 3 years. Under such regimes, emergence and early growth of ginger are inherently slow and considerable time elapses between sowing and the development of foliage cover (Lee et al., 1981). In addition, ginger normally propagates with a low proliferation rate (about 10–15 buds from one plant each year) by its rhizome, and is easily infected by soil-born
* Corresponding author. Tel.: +86 28 85248450. E-mail addresses: hanchunmei@tom.com (C.-M. Han), pankw@cib.ac.cn (K.-W. Pan). 0304-4238/$ – see front matter # 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.scienta.2008.01.005

pathogens such as bacterial wilt (Pseudomonas solanacearum), soft rot (Pythium aphanidermatum), and nematodes (Meloidogyne spp.), which cause heavy losses in yields (Guo and Zhang, 2005). Autotoxicity is a type of intraspeci?c allelopathy, where a plant species inhibits the growth of its own kind through the release of toxic chemicals into the environment (Singh et al., 1999). This phenomenon has been demonstrated in a number of crop plants such as annual crops like wheat, rice, maize, mungbean grown in monocultures, forage crops like alfalfa and clover, and oil crops such as sun?ower and rapeseed, and others like asparagus, sugarbeet, cucumber, carrot, coriander, cumin and fennel. In most cases, it is related to the repeated sowing of monocultures leading to soil sickness (Singh et al., 1999). Autotoxicity may be one reason for lower production rates of successive crops of ginger. In China, the Shandong Province, Zhejiang Province, Guangdong Province and Sichuan Province are the main production areas of ginger. The use of other crops may alleviate the problem of autotoxicity caused by repeatedly planting ginger monocultures. Soybeans (Glycine max (L.) Merr.) and chives (Allium schoenoprasum L.) are two promising crops for intercropping with ginger. Allelopathy is de?ned as any direct or indirect positive or negative effect of one plant on the other (including the

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microbes) through the release of chemicals into the environment (Rice, 1984). It plays a signi?cant role in agroecosystems, and affects the growth, quality and quantity of the produce (Kohli et al., 1998; Singh et al., 2001). A number of plant species have been reported to have an allelopathic effect on other plant species (Mallik, 1987; Martin and Smith, 1994; Kato-Noguchi, 2003; Jefferson and Pennacchio, 2003; Oueslati, 2003; Djurdjevic et al., 2004). Allelochemicals produced by one crop species can in?uence the growth, productivity, and yield of other crops or the same crop (Batish et al., 2001). The avoidance of allelopathic effects between crops, or the exploitation of bene?cial interactions in a rotation or a mixed cropping system may have direct bearing on the crop yield (Rizvi et al., 1992). In the present study, the allelopathic effects of ginger rhizome, stem and leaf aqueous extracts were examined to determine if inhibitory or stimulatory effects of ginger extracts in?uence seed germination and seedling growth of two crops (soybean and chive) that are commonly intercropped with ginger. 2. Materials and methods

as a control treatment. All Petri dishes were placed in a dark room at 25 8C. Treatments were arranged in a completely randomized design with three replications. Germination was determined by counting the number of germinated seeds at 24-h intervals over a 6-d (soybean) and 17-d (chive) period and expressed as total percent germination. Germination was deemed to occur only after the radicle had protruded beyond the seed coat by at least 1 mm. Radicle and hypocotyl lengths of soybean and chive seedlings were measured 6 d and 17 d after germination, respectively. After measuring the radicle and hypocotyl lengths, the dry weights of seedlings were determined by drying the plant material in an oven at 60 8C for 24 h prior to weighing. The inhibitory or stimulatory percent was calculated using the following equation given by Chung et al. (2001): Inhibition??? or stimulation??? percentage ?%?   extracts ? control ? 100: ? control 2.4. Water uptake

2.1. Location Ginger plants were collected from ?elds (29815 N, 1038470 E, 510 m altitude) in Sichuan Province, Leshan City, China, in August 2005. The experiment was carried out at the ecological center, Chengdu Institute of Biology, Chinese Academies of Sciences, from July to October 2006. 2.2. Preparation of extracts Fresh ginger plants were separated into leaves, stems and rhizome. The stems and leaves were chopped into 1 cm long pieces and rhizomes were chopped into 0.5 cm thick slices. The components were then oven dried at 60 8C for 5 days. Eighty grams of dried rhizomes, stems and leaves were respectively extracted by soaking in 1 l deionized water at 25 8C for 24 h in a shaker to give a concentration of 80 g dry tissue l?1 (g l?1). The extracts were respectively ?ltered through four layers of cheesecloth to remove the ?ber debris, and centrifuged at 3000 rpm for 4 h (Chon et al., 2002). The supernatant was ?ltered again using a 0.2-mm ?lterware unit. Fresh stock extracts were kept in a refrigerator at 2 8C until used. 2.3. Seed bioassay Stock extracts (rhizome, stem and leaf) were diluted with sterile distilled water to give ?nal concentrations of 0, 10, 20, 40 and 80 g l?1. Seed germination tests were conducted for each extract as follows: 30 soybean and chive seeds were surface sterilized with 5.25% (w/v) sodium hypochlorite solution for 15 min, rinsed three times with distilled water and were evenly placed on two-layer ?lter paper in sterilized 9-cm Petri dishes. Due to differences in the size of soybean and chive seed, 15 and 5 ml of extract solution were added to Petri dish containing soybean and chive seeds, respectively. Distilled water was used
0

Approximately 1-g samples (W1, the original seed weight) of soybean and chive seeds were separately soaked for 4, 8, 12, 16 and 20 h in the aqueous extracts. Distilled water was used as the control treatment. At 4-h intervals, seeds were taken from the solution, blotted for 3 h between two folds of ?lter paper, and weighed (W2, the ?nal seed weight). This is based on the method given by Turk and Tawaha (2003). Water uptake percentage is expressed as follows: Water uptake ?%? ? W2 ? W1 ? 100: W1

2.5. Assay of lipase activity of soybean and chive seed After measuring water uptake, the seeds were shelled, grounded and extracted for 1 h with 0.1 M Tris–HCl buffer (pH 7.0) at 25 8C. After centrifugation, the soluble proteins in the supernatant were fractionated and precipitated with 50% (v/v) and 80% (v/v) ethanol at 4 8C. The precipitates obtained were dissolved in 0.1 M Tris–HCl buffer (pH 7.0). To prepare an acetone powder, the germinated seeds were ground in ice-cold acetone and then washed several times with the cold acetone (Ncube et al., 1995). Lipase (glycerol ester hydrolase, E.C. 3.1.1.3) activity was assayed using a modi?cation of the titrimetric method of Khor et al. (1986). The assay mixture contained 5 g of substrate, 2.5 ml of hexane to solubilize the oil, and 0.5 g of the crude enzyme. The mixture was incubated at 30 8C for a period of 1 h with continuous stirring, using a magnetic stirrer. At the end of the incubation, 25 ml of acetone– ethanol (1:1, v/v) were added to stop the reaction and to extract the free fatty acids (FFAs) liberated. The FFAs in the mixture were then estimated by direct titration with 0.01 M NaOH using phenolphthalein as an indicator of lipase activity. Lipase activity was expressed as the percent FFAs liberated after 1 h

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Table 1 Effects of ginger aqueous extracts on seed germination and seedling growth of soybean Ginger parts Rhizome Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Germination (%) 98 ? 1.0 97 ? 1.9 100 ? 0.0 96 ? 2.2 91 ? 5.6 ns ns ns ns Hypocotyl length (cm) 5.63 ? 0.26 5.42 ? 0.31 ns 4.99 ? 0.13 ns 4.15 ? 0.12** 3.22 ? 0.34** 4.43 ? 0.17** 3.07 ? 0.20** 0.60 ? 0.31** 0.00 ? 0.00** 4.93 ? 0.28 ns 5.03 ? 0.25 ns 3.55 ? 0.42* 1.31 ? 0.94** Radicle length (cm) 5.35 ? 0.62 5.95 ? 4.07 ns 4.67 ? 2.21 ns 4.38 ? 3.70 ns 2.55 ? 1.08** 3.18 ? 0.20** 0.55 ? 0.31** 0.00 ? 0.00** 0.00 ? 0.00** 6.68 ? 0.20 ns 6.63 ? 0.63 ns 3.25 ? 0.74 ns 0.43 ? 0.43** Dry weight per seedling (mg) 23.4 ? 0.5 24.4 ? 1.2 ns 23.7 ? 1.2 ns 23.5 ? 1.5 ns 16.3 ? 1.1** 20.2 ? 0.9 ns 8.0 ? 3.1** 3.0 ? 1.6** 0.0 ? 0.0** 25.2 ? 0.4 ns 24.7 ? 2.0 ns 19.6 ? 2.5 ns 6.1 ? 6.1**

Stem

96 ? 1.1 ns 46 ? 8.7** 2 ? 1.1** 0 ? 0.0** 93 ? 1.9 ns 98 ? 2.2 ns 83 ? 3.8* 7 ? 5.1**

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

incubation (Wetter, 1957). Corrections were made for endogenous fatty acid production (assay mixture without substrate) and nonenzymatic fatty acid production (assay mixture without enzyme preparation). 2.6. Experimental design and statistical analysis Germination and seedling growth bioassays were calculated in a complete randomized design with three replications. The data were subjected to one-way analysis of variance, and treatment means separated from the control at P < 0.05 or 0.01 applying post hoc Dunnett’s test. Statistical analysis was done with SPSS 11.0 for Windows statistical software package. 3. Results 3.1. Effect on seed germination and seedling growth of soybean and chive The allelopathic potential of aqueous extracts of rhizome, stem and leaf from ginger was tested on the germination, radicle

and hypocotyl lengths, and dry weight per seedling of two test species (Tables 1 and 2). The germination of the control was 98%. Application of 20 g l?1 aqueous extract of rhizome had a stimulatory but not signi?cant effect (2.2%) on soybean seed germination (Table 1). Increased concentration of ginger extracts from all plant parts inhibited germination. The degree of inhibition increased with increasing extract concentration. At the highest extract concentration (80 g l?1), stem and leaf aqueous extracts signi?cantly inhibited seed germination compared with the control. Stem extracts were mostly inhibitory at 20 g l?1 or greater, while the rhizome extracts were the least inhibitory. Stem extract inhibited germination by 53.4%, 97.8% and 100.0% at the 20, 40 and 80 g l?1, respectively. All aqueous extracts at all concentrations inhibited hypocotyl length compared with the control and the degree of inhibition increased with increasing extract concentration. The highest concentration (80 g l?1) of stem extract resulted in no hypocotyl development. Adverse effect of stem extract on radicle length and dry weight was similar to that of the hypocotyl (Table 1). The effects of rhizome and leaf extracts on radicle length and dry

Table 2 Effects of ginger aqueous extracts on seed germination and seedling growth of chive Ginger parts Rhizome Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Germination (%) 81 ? 3.0 74 ? 4 ns 77 ? 6 ns 62 ? 5 ns 44 ? 7** 73 ? 7 ns 50 ? 3** 36 ? 3** 6 ? 4** 73 ? 2 ns 66 ? 2 ns 32 ? 8** 31 ? 10** Hypocotyl length (cm) 5.38 ? 0.21 4.11 ? 0.26* 3.28 ? 0.55** 2.16 ? 0.25** 0.06 ? 0.06** 2.93 ? 0.17** 0.27 ? 0.14** 0.00 ? 0.0** 0.00 ? 0.0** 3.50 ? 0.29** 2.51 ? 0.22** 0.68 ? 0.17** 0.00 ? 0.0** Radicle length (cm) 1.69 ? 0.19 0.95 ? 0.06** 0.78 ? 0.19** 0.43 ? 0.03** 0.38 ? 0.08** 0.62 ? 0.09** 0.40 ? 0.06** 0.29 ? 0.01** 0.17 ? 0.03** 0.90 ? 0.08** 0.63 ? 0.16** 0.48 ? 0.02** 0.43 ? 0.03** Dry weight per seedling (mg) 0.79 ? 0.02 0.78 ? 0.06 ns 0.75 ? 0.22 ns 0.56 ? 0.07 ns 0.05 ? 0.05** 0.63 ? 0.20 ns 0.12 ? 0.06** 0.00 ? 0.00** 0.00 ? 0.00** 0.76 ? 0.05 ns 0.67 ? 0.17 ns 0.28 ? 0.06** 0.00 ? 0.00**

Stem

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

C.-M. Han et al. / Scientia Horticulturae 116 (2008) 330–336 Table 3 Effects of ginger aqueous extracts on water uptake of soybean seeds at different imbibition periods Ginger parts Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Water uptake of soybean (%), by imbibition period (h) 4 93.8 91.9 ns 91.5 ns 83.3** 76.2** 84.9* 84.1* 79.4** 73.5** 89.0 ns 86.8 ns 83.4* 74.7** 8 116.4 116.3 ns 116.0 ns 114.2 ns 106.1** 114.4 ns 112.4 ns 107.1** 102.2** 116.3 ns 115.4 ns 112.0 ns 106.7** 12 118.8 117.8 ns 117.1 ns 116.8 ns 111.7** 116.5 ns 114.0 ns 110.2** 109.5** 117.5 ns 117.0 ns 116.7 ns 111.7* 16 120.6 119.4 ns 119.1 ns 117.9 ns 114.0* 119.7 ns 117.4 ns 113.9* 112.5** 120.4 ns 119.4 ns 117.8 ns 113.6** 20

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Rhizome

121.5 120.7 ns 120.7 ns 118.6 ns 115.4** 120.1 ns 118.9 ns 114.8** 113.7** 121.2 ns 119.9 ns 118.1 ns 114.7**

Stem

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

weight both showed a tendency of being stimulatory at lower concentrations and becoming inhibitory at higher concentrations. Maximum radicle length and dry weight were recorded at 10 g l?1 of leaf aqueous extract, and their values were 24.9% and 7.7% higher than those of the control, respectively. The radicle length was more sensitive to all types of extract used in comparison to the other plant parameters measured. The effect of ginger aqueous extracts on germination, radicle length, hypocotyl length and dry weight of chive was stronger than that of soybean (Table 2). All plant parameters measured were reduced with the increase in different extract concentrations and all of them reached minimum values with application of 80 g l?1 stem extract concentration. Between the two test species, a greater inhibitory effect was observed on chive compared with soybean (Tables 1 and 2). 3.2. Effect on the rate of water uptake of soybean and chive A similar trend of changes was observed with regard to water uptake of soybean and chive. Increasing the concentrations of

aqueous rhizome, stem and leaf extracts inhibited water uptake to different extent by germinating soybean and chive seeds at different imbibition periods. The greatest inhibition of water uptake by soybean and chive seeds occurred when seeds were imbibed for 4 h in 80 g l?1 of stem extract (Tables 3 and 4). The percent of water uptake was 73.5% for soybean and 29.2% for chive which decreased by 21.6% and 28.6%, respectively, compared with the control. The percentage of water uptake of soybean seed was twice as much as that of chive seed under the same aqueous extract concentration. Moreover, under the same concentration, the percentage of water uptake increased by prolonging the imbibition period. 3.3. Effect on the lipase activities of soybean and chive Lipase activity was inhibited in a dose-dependent manner (Tables 5 and 6). Under the same extract concentration, lipase activity increased by prolonging the imbibition period. However, for soybean seeds, lipase activity was not signi?cantly inhibited at all concentrations of aqueous extracts

Table 4 Effects of ginger aqueous extracts on water uptake of chive seeds at different imbibition periods Ginger parts Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Water uptake of chive (%), by imbibition period (h) 4 40.9 39.7 ns 38.1* 36.2** 35.1** 36.4** 35.2** 32.9** 29.2** 39.0 ns 37.6** 35.6** 34.8** 8 53.5 53.0 ns 51.8 ns 49.9 ns 47.1* 49.9 ns 46.0** 44.2** 38.6** 52.5 ns 50.2* 49.6* 46.4** 12 56.7 56.3 ns 53.8 ns 52.0* 50.2** 51.2** 48.8** 45.4** 40.8** 55.6 ns 52.8* 51.6** 48.7** 16 58.6 57.4 ns 55.3 ns 53.6* 51.6** 52.3** 49.5** 48.1** 39.6** 56.4 ns 53.6* 52.6** 50.2** 20 59.7 58.2 ns 56.2* 54.3** 52.3** 53.1** 50.2** 48.7** 39.6** 56.9* 54.3** 53.1** 50.9**

Rhizome

Stem

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

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Table 5 Effects of ginger aqueous extracts on lipase activity of soybean seeds at different imbibition periods Ginger parts Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Lipase activity of soybean (% FFAs), by imbibition period (h) 4 1.47 1.42 1.28 1.23 0.90 1.30 1.25 1.16 0.76 1.33 1.26 1.18 0.87 ns ns ns ns ns ns ns ns ns ns ns ns 8 2.26 2.10 2.00 1.85 1.69 ns ns ns ns 12 2.77 2.74 2.59 2.46 2.33 2.64 2.56 2.40 2.30 2.66 2.57 2.41 2.32 ns ns ns ns ns ns ns ns ns ns ns ns 16 3.45 3.40 3.25 2.99 2.89 ns ns ns ns 20 4.48 4.23 ns 3.98 ns 3.90* 3.63** 4.06 ns 3.92* 3.73** 3.53** 4.11 ns 3.97 ns 3.83* 3.59**

Rhizome

Stem

2.03 ns 1.90 ns 1.77* 1.57** 2.07 ns 1.97 ns 1.80* 1.66**

3.28 ns 3.08 ns 2.96 ns 2.48** 3.30 ns 3.20 ns 2.98 ns 2.87*

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

when seeds were imbibed for 4 h and 12 h. It was however signi?cantly inhibited when imbibing the seeds in stem and leaf extracts at concentrations of 40 or 80 g l?1 for a period of 8 h and 16 h. When imbibing the seeds for a period of 20 h, lipase activity was signi?cantly inhibited at 20 g l?1 or greater of all aqueous extracts (Table 5). The same trend existed in lipase activity of chive seeds. Lipase activity of chive seeds was however signi?cantly inhibited at the shortest imbibition time of 4 h (Table 6). 4. Discussion The results indicate that aqueous extracts from different parts of ginger show a phytotoxic in?uence on soybean and chive. The phytotoxic effect was differential and tissue speci?c: stem > leaf > rhizome. The degree of inhibition was largely dependent on the concentration of the extracts being tested. The residue (mainly stems and leaves) of the ginger plant exhibits stronger phytotoxicity. These results indicate that ginger

residues release allelopathic substances which accumulate in bioactive concentrations and adversely affect seed germination, seedling growth, water uptake and lipase activity of soybean and chive. Extracts from ginger rhizome, stem and leaf solutions showed inhibitory effects on seed germination of two test species. The degree of inhibition increased with increasing extract concentration. At the highest extract concentration (80 g l?1), all aqueous extracts (except for rhizome extract for soybean) signi?cantly inhibited seed germination of two test species compared with the control (Tables 1 and 2). This ?nding is congruent with the results of Chung and Miller (1995) who found that the degree of inhibition increased with increasing extract concentration. Stem extracts exhibited the greatest inhibition at all concentrations, while rhizome extracts was the least inhibitory. The results found in this study are inconsistent with those of Turk and Tawaha (2003), who reported that leaf extracts of black mustard (Brassica nigra L.) exhibited the greatest inhibition, while stem extracts were the least inhibitory

Table 6 Effects of ginger aqueous extracts on lipase activity of chive seeds at different imbibition periods Ginger parts Extract concentration (g l?1) 0 10 20 40 80 10 20 40 80 10 20 40 80 Lipase activity of chive (% FFAs), by imbibition period (h) 4 0.75 0.71 ns 0.68* 0.52** 0.43** 0.65** 0.53** 0.42** 0.31** 0.69** 0.60** 0.49** 0.41** 8 1.13 1.09 ns 0.98 ns 0.87** 0.71** 0.99** 0.78** 0.64** 0.58** 1.01** 0.92** 0.81** 0.70** 12 2.25 2.13* 2.06** 1.95** 1.83** 1.94** 1.70** 1.59** 1.47** 2.08* 1.94** 1.85** 1.68** 16 3.08 2.92* 2.81** 2.76** 2.65** 2.71** 2.53** 2.29** 2.04** 2.84** 2.69** 2.54** 2.38** 20 3.83 3.74 ns 3.61** 3.43** 3.27** 3.52** 3.31** 3.19** 3.07** 3.65** 3.49** 3.31** 3.19**

Rhizome

Stem

Leaf

ns: not signi?cant; (*) and (**) represent signi?cant difference over control at P < 0.05 and P < 0.01, respectively.

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(the test species is wild oat (Avena fatua L.)). However, it has been reported by Turk and Tawaha (2002) that leaf extracts of black mustard is the most inhibitory, while root extract is the least inhibitory (the test species is lentil (Polygala tatarinowii Regel.)). The reason for the differences between the two studies may be due to the fact that two different test species were used. A number of studies have suggested that plant residues (especially weed species) affect the growth and development of other plants including crops by releasing allelochemicals into the immediate soil environment (Singh et al., 2003a,b, 2005; Batish et al., 2006a,b). Hegde and Miller (1992) reported the adverse effects of phytotoxins from crop residues on the seedling growth of succeeding crops. Recently, Batish et al. (2007) reported that nettle-leaved goosefoot (Chenopodium murale) roots and their exudates exerted allelopathic effects on wheat by releasing water-soluble phenolic acids as putative allelochemicals in soil. The present study demonstrated that all ginger aqueous extracts, especially stem and leaf (ginger residue) extracts inhibited seedling growth of two test species (Tables 1 and 2). The radicle length was found to be more sensitive to all types of ginger aqueous extract in comparison to the other parameters measured. These results are in agreement with those of Turk and Tawaha (2002). Between the two test species, a greater inhibitory effect was observed on chive than on soybean, which suggests species speci?city. Water uptake of two test species was signi?cantly inhibited with increasing extract concentration (Tables 3 and 4), which was similar to results of Turk and Tawaha (2003) who reported that increasing the concentration of black mustard aqueous leaf extracts signi?cantly inhibited the water uptake of germinating wild oat seeds. There may be an indirect association between lower seed germination and allelopathic inhibition of water uptake and lipase activity. Water uptake of soybean seed was obviously faster than that of chive seed under the same aqueous extract concentration, which might attribute to the size of soybean and chive seeds. It may also be related to the physical properties of the test seeds. Soybean seeds absorbed water faster than chive seeds, probably because of a thinner seed coat, which can be more easily penetrated by water (Bewley and Black, 1985). However, the size of test seeds may be the limiting factor for water uptake in this condition. Moreover, the percentage of water uptake increased by prolonging the imbibition period under the same concentration, which might attribute to meeting the need for seed germination. Several enzymes like proteases, lipases and a-amylases play an important role during seed germination. Many enzymatic functions are inhibited by the presence of allelochemicals (Turk ?as and Tawaha, 2002; Rice, 1984). Kato-Noguchi and Mac? (2005) had reported that the germination of lettuce (Lactuca sativa L. cv. Grand Rapids) seeds treated by 6-methoxy-2benzoxazolinone (MBOA) was positively correlated with the activity of a-amylase. In soybean and chive seeds, lipases function to mobilize storage triglycerides through hydrolysis to fatty acids during early stages of germination. The fatty acids released by the lipases are channeled into energy producing pathways thereby providing energy for the growing embryo and seedling (Staubmann et al., 1999).

Lipase activity of both test species showed inhibitory tendency with increasing concentration of ginger aqueous extracts (Tables 5 and 6), which was consistent with that of water uptake. This ?nding is supported by Moreno et al. (2003) who found that grape seed extract inhibited lipase activity in a dose-dependent manner. There may be an indirect association between lower lipase activity and allelopathic inhibition of water uptake. To a large extent, lipase activity is primarily related to water uptake by the seeds (Turk and Tawaha, 2003). Lipase activity of soybean seed was signi?cantly higher than that of chive seed under the same aqueous extract concentration (Tables 5 and 6). This result might attribute to the size of test plant seeds. Soybean seed is bigger than that of chive and its water uptake is relatively faster than that of chive seed, which might lead to higher lipase activity. In the present investigation, no attempt was made to identify the allelochemicals produced by ginger and to conduct bioassays under realistic soil conditions. However, the present laboratory bioassays indicate the presence of some watersoluble phytotoxins in ginger extracts that leach from the debris into the water. Further work is needed to specify and verify the allelochemicals produced by ginger and validate ginger allelopathy under actual ?eld conditions in its habitat. Such studies using soil to simulate natural environmental conditions carry great ecological signi?cance. 5. Conclusion Ginger residues (mainly leaves and stems) adversely affect seed germination and seedling growth of two crops (soybean and chive) that are commonly intercropped with ginger. Therefore, ginger must be considered as an allelopathic species posing risk in a rotation or an intercropping or mixed cropping system. With a view to alleviate its adverse effects on intercropping or subsequent crops, farmers should remove residues of ginger from the agricultural land. Acknowledgements The research was supported by the cooperation program of Chinese Academy of Sciences and Leshan City (No. 2005BA807B09LA06) and the China National Key program of ‘‘ Tenth Five-year Plan’’ (No. 2004BA606-05-03). References
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