Effects of indigenous arbuscular mycorrhizal fungi in paddy fields on

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Effects of indigenous arbuscular mycorrhizal fungi in paddy fields on rice growth and N, P, K nutrition under different water regimes Mohammad Zakaria Solaiman

a b

& Hiroshi Hirata

a c

a

Laboratory of Plant Nutrition, Department of Biological Production, Faculty of Agriculture, Tokyo University of Agriculture and Technology, Fuchu, 183, Japan b

Department of Soil Science, Bangladesh Agricultural University, Mymensingh, 2202, Bangladesh c

Laboratory of Plant Nutrition, Department of Biological Production, Faculty of Agriculture, Tokyo University of Agriculture and Technology, Fuchu, 183, Japan Available online: 04 Jan 2012

To cite this article: Mohammad Zakaria Solaiman & Hiroshi Hirata (1995): Effects of indigenous arbuscular mycorrhizal fungi in paddy fields on rice growth and N, P, K nutrition under different water regimes, Soil Science and Plant Nutrition, 41:3, 505-514 To link to this article: http://dx.doi.org/10.1080/00380768.1995.10419612

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Soil ScL Plant Nutr., 41 (3), 505-514, 1995

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Effects of Indigenous Arbuscular Mycorrhizal Fungi in Paddy Fields on Rice Growth and N, P, K Nutrition under Different Water Regimes

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M o h a m m a d Z a k a r i a S o l a i m a n 1 and H i r o s h i H i r a t a 2 Laboratory of Plant Nutrition, Department of Biological Production, Faculty of Agriculture, Tokyo University of Agriculture and Technology, Fuchu, 183 Japan Received May 16, 1994; accepted in revised form March 2, 1995 A pot experiment was conducted under two soil conditions, namely (i) Original paddy soil and (ii) Diluted soil (paddy soil diluted with andosol subsoil; Paddy soil : Andosol s u b s o i l = l : 4) to i n v e s t i g a t e the effects of arbuseular m y c o r r h i z a l fungi (AMF) and w a t e r regimes on dry m a t t e r production, g r a i n yield, A M F colonization, A M F sporulation, and mineral n u t r i t i o n of wetland rice (Or!tza sativa L.). P l a n t s were e i t h e r inoculated ( + A M F ) or not ( - A M F ) under four w a t e r regimes, namely (i) Flooded up to m a t u r i t y (F); (ii) Non-flooded (kept at 60% moisture of maximum w a t e r holding capacity) up to m a t u r i t y (NF); (iii) Flooded for 3 weeks a f t e r sowing and t h e n non-flooded up to m a t u r i t y (F-NF); and (iv) Non-flooded f o r 3 weeks a f t e r sowing and t h e n flooded up to m a t u r i t y (NF-F). T h e r e were 2 X 2 • t r e a t m e n t combinations. Generally, plants g r o w n in paddy soil showed a h i g h e r plant dry m a t t e r c o n t e n t and h i g h e r N, P, K c o n c e n t r a t i o n s t h a n in diluted soil. Grain yield was h i g h e r and grain f o r m a t i o n was improved under flooded (F and N F - F ) t h a n under non-flooded conditions (NF and F-NF) and g r a i n f o r m a t i o n was r e t a r d e d in the l a t t e r resulting in a low h a r v e s t index especially in inoculated plants. Mycorrhiza] t r e a t m e n t resulted in a decrease in the a m o u n t of shoot dry m a t t e r under flooded conditions and also in the production of unhulled grains under nonflooded conditions. T h o u g h m y c o r r h i z a l colonization was highest under nonflooded conditions, a 2-12% colonization level was observed even under flooded conditions at 60 d a f t e r t r a n s p l a n t i n g , and 3% remained in the N F - F t r e a t m e n t whereas no colonization occurred under continuously flooded conditions (F) at the m a t u r a t i o n stage. AM establishment before flooding had a positive impact on A M F colonization and sporulation in wetland rice cultivation at the vegetative stage. N and P c o n c e n t r a t i o n s in unhulled grain were significantly increased due to A M F inoculation whereas A M F significantly decreased shoot N c o n c e n t r a t i o n and did not affect shoot P c o n c e n t r a t i o n at the m a t u r a t i o n stage. These findings suggest t h a t A M F may accelerate N and P t r a n s f e r f r o m shoots a n d / o r soils to rice grains even under flooded conditions along with the t e n d e n c y to increase t h e h a r v e s t index. K e y Words: arbuscular m y c o r r h i z a l fungi, indigenous, soil properties, w a t e r regimes, wetland rice. :Present address: Department of Soil Science, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh. 2 To whom correspondence should be addressed.

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The endomycorrhizal association is a finely balanced complex system involving a three-way interaction among plant, fungal endophyte, and soil (Hayman 1983). Analysis of both this association and the changes in the population of infective arbuscular mycorrhizal fungi (AMF) in an agricultural ecosystem is necessary for proper management of beneficial symbiosis. Cultivation under different cultural practices and soil conditions is likely to affect the A M F population in the soil. Plants associated with A M F show a range of responses, from mutualistic to parasitic. The responses observed may depend on the soil moisture, availability of nutrients, pH, A M F species, host plant species, type of host root system, growth stage of host plant and irradiation, among other factors (Dhillion and Ampornpan 1992). In most studies the effects of AM fungi on plant growth and nutrient uptake, notably P (Hayman 1983; Koide 1991) have been examined and studies investigating the nutrient content of AM and non-AM plants have revealed inconsistent results (Harley and Smith 1983). However, the promotion of plant growth associated with mycorrhizal association has been attributed to improved nutrient status (Hayman 1983). Growth depression associated with A M F colonization has been ascribed to host-fungus competition for carbohydrates (Buwalda and G o h 1982) or to interactions between soil organisms and AM fungi (Bagyaraj 1984). In only a few recent studies the mycorrhizal nature of rice has been investigated (Ilag et al. 1987; Brown et al. 1988; Dhillion and Ampornpan 1990, 1992). Although the AM associations are generally less frequent in wetland rice cultivation than upland cultivation, some reports indicated that wetland rice plants in submerged fields were highly colonized (Ilag et al. 1987; Sivaprasad et al. 1990; Secilia and Bagyaraj 1992). Sivaprasad et al. (1990) reported that grain and straw yields in rice plants inoculated with Glornus fasciculatum were significantly higher than in non-mycorrhizal ones and concluded that A M F inoculation by the dry nursery method may enhance AM benefits in lowland (wetland) rice cultivation. On the other hand, Dhillion and Ampornpan (1990) reported a significant reduction in shoot and root growth in mycorrhizal plants compared with non-mycorrhizal plants when the plants received supplements of P and N. The objective of this experiment was to investigate the effects of indigenous AM fungi in paddy fields on the growth and N, P, K nutrition of rice under different water regimes and soil properties. MATERIALS AND METHODS Sites a n d soils. The soils used in this experiment included paddy soil (alluvial loamy soil; collected from the Tokyo University of Agriculture and Technology farm, Fuchuhonmachi, Tokyo, Japan) and diluted soil. The diluted soil consisted of a mixture of paddy soil and of a red Andosol subsoil in the ratio of 1 : 4. The subsoil was collected from the Tokyo University of Agriculture and Technology farm field, Saiwai-cho, Tokyo, Japan. Both soils were sterilized by 15 MGray gamma irradiation. Some of the chemical properties of the soils used in this experiment are shown in Table I. The total N and C contents of the soils were determined with a NC analyzer (Sumitomo Chemical, S U M I G R A P H NC 80). Other soil properties were determined based on some standard methods (Page et al. 1990). I n o e u l u m . Fresh soil collected from the paddy field continuously cultivated with rice was used as medium and source of indigenous A M F inoculum. Inoculum soil which contained 335 spores on an average per 100 g fresh soil was measured by using a 53 a m mesh. The inoculated fungi were mainly of Glomus sp. For the mycorrhizal treatment, 100 g fresh non-sterilized soil was inoculated as layer application at 5 cm depth and for the non-mycorrhizal treatment an equivalent amount of gamma ray sterilized soil was added to the pot.

Indigenous Arbuscular Mycorrhizas in Rice Cultivation

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Table 1. Some chemical properties of soils used. Properties Paddy soil Andosol subsoil pH (H~O) 6.0 5.4 Total carbon (g kg -1) 44.5 12.6 Total nitrogen (g kg -1) 3.83 0.89 Bray 2P (mg kg-1) 171.2 1.37 Total P (mg kg-1) 1307.5 50.85 Exchangeable K (mg kg -1) 370 120

507 Diluted soil 5.7 18.1 1.37 30.39 454.2 220

Treatments. Plants were inoculated ( § or not ( - - A M F ) under two soil conditions namely (i) Paddy soil (PS), and (ii) Diluted soil (DS) under four water regimes, (i) Flooded up to maturity (F); (ii) Non-flooded up to maturity (NF); (iii) Flooded for 3 weeks after sowing and then non-flooded up to maturity (F-NF); and (iv) Non-flooded for 3 weeks after sowing and then flooded up to maturity (NF-F). Therefore, there were 2 x 2 • 4 = 16 treatment combinations. This study was conducted as a 2 • 2 1 5 factorial experiment in a completely randomized block design with three replications. G r o w t h .~f p l a n t . A cultivar of wetland rice, Oryza sativa L. var. Japonica cv. N i p p o n b a r e was used. The plants were grown in 5 X 10 -3 a Wagner pots under glass house conditions. The plants were watered daily with deionized water and the flooding level was maintained at about 3 cm from the soil surface. Non-flooded pots were kept at 60% moisture level of maximum water holding capacity during the growth period. N, P, and K fertilizers were applied at the rate of 700, 0, and 500 mg per pot, respectively. The N from urea was applied in 3 splits: 300 mg N as basal dressing, 200 mg at 60 d (maximum tillering stage), and the rest (200 mg) at 90 d after sowing (heading stage). K from muriate of potash (KC1) was applied only as basal dressing. Six pregerminated rice seeds were sown in each pot and then thinned to two per pot. The period of growth (June 1992 to November 1992) was 140 d for the flooded conditions (F; N F - F ) and 155 d for the non-flooded conditions (NF; F-NF). S a m p l i n g . Sampling was performed at two growth stages, one at the m a x i m u m tillering stage (60 d after sowing) and the other at harvest (140/155 d after sowing). A m o n g the data from the first sampling (60 d after sowing) only the A M F colonization and the no. of spores per pot were presented in this paper. M y e o r r h i z a l a s s a y s . Spores of A M F in the soil were estimated by the wet sieving and decantation method described by Daniels and Skipper (1984). At both sampling times, a 1.0 g subsample of the roots was excised from each plant, to assess the percentage of A M F colonization. The root samples were cleared in K O H (100 g L -1) solution at 90~ on a hot plate for 1 h and stained with trypan blue (0.5 g L -1) in lactoglycerol ( K o r m a n i k and McGraw 1984) at 90~ for 30 min. Percentage colonization of host plant roots was estimated by visual observations of stained root segments mounted in lactoglycerol by the grid-line intercept method (Giovanetti and Mosse 1980). P l a n t tissue a n a l y s i s . All the tissue samples were oven-dried at 80~ for 48 h and weighed. Oven-dried grains, shoots (leaves and stems), and roots were analyzed for N, P, and K contents. The plant P concentration was determined by the vanadomolybdate blue method; Nitrogen by the I n d o p h e n o l colorimetric method and Potassium by flame photometry (Page et al. 1990). S t a t i s t i c a l a n a l y s i s . Three-way analysis of variance ( 2 • 2 1 5 combining 4 water regimes into two (because the trend of data among them was similar) namely flooded (F and N F - F ) and non-flooded (NF and F - N F ) was performed to determine the main and interac-

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tion effects of different factors on AM formation, sporulation, plant growth, nutrient (N, P, K) concentrations of rice plants. Duncan's multiple range test (DMRT) or Least significant difference (LSD) test for mean separation was used to determine significant differences among treatment means. Statistical significance was accepted at p < 0.05 level of significance.

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RESULTS

Dry matter production As shown in Fig. i, the plants under non-flooded conditions (NF and F - N F ) in the paddy soil almost failed to produce grains compared to the plants under flooded conditions whereas the differences between flooded and non-flooded conditions were negligible in diluted soil. Generally, under flooded conditions (F and N F - F ) of both soils (PS and DS), A M F inoculation resulted in the decrease of dry matter production of shoots though the dry weight of unhulled grains and roots did not differ between the inoculated and non-inoculated treatments. On the other hand, under non-flooded conditions (NF and F-NF) opposite phenomena were observed, especially in N F under both soil conditions.

fig. 1. Effect of soil properties and water regimes on (a) unhulled grain, (b) shoot (leaves and stems), and (c) root dry matter of mycorrhizal and nonmycorrhizal wetland rice at maturation stage (140/155 DAS). F=Flooded up to maturity; NF-F=Nonflooded 3 weeks and then flooded up to maturity; NF=Non-flooded up to maturity; F-NF= Flooded 3 weeks and then non-flooded up to maturity; --AMF=Not inoculated; +AMF=Inoculated; DAS=Days after sowing.

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Indigenous Arbuscular Mycorrhizas in Rice Cultivation

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Grain f o r m a t i o n and grain h a r v e s t index At the maturation stage, m a x i m u m number of grains ripened and plants became yellowish under flooded conditions which under non-flooded conditions the plants remained green and almost failed to produce filled grains especially in the original paddy soil. Grain formation under flooded conditions was significantly improved compared with non-flooded conditions (Fig. 2a). The differences in grain formation between flooded and non-flooded conditions were less appreciable in diluted paddy soil with andosol than in paddy soil. The grain harvest index of wetland rice in the mycorrhizal treatment tended to be higher than in the non-mycorrhizal one under flooded conditions whereas under nonflooded conditions reverse effect was observed in both soils (Fig. 2b and Table 3).

AMF colonization The A M F colonization of root was affected by the water regime. The A M F colonization was highest under non-flooded conditions (NF), where as a 2-12% infection level was observed under flooded conditions (F; N F - F ) at the maximum tillering stage (60 d after sowing) irrespective of differences in soil properties but up to the maturation stage 3% infection persisted in the N F - F water regime treatment under both soil conditions (Fig. 3a). Under non-flooded conditions in paddy soil A M F colonization was higher than in diluted soil. The A M F colonization of roots decreased along with the growth stages irrespective of the differences in water regimes and soil conditions.

Fig. 2. Effect of soil properties and water regimes on (a) grain formation and (b) harvest index of mycorrhizal and non-mycorrhizal wetland rice at maturation stage (140/155 DAS). See footnote to Fig. 1. Grain formation=(Filled grain)/(Filled grain+ Empty grain)• Harvest index= (Grain)/(Grain+Leaves & Stems) • 100.

Fig. 3. Effect of soil properties and water regimes on (a)AMF colonization and (b)AMF sporulation of mycorrhizal wetland rice at two different growth stages. See footnote to Fig. 1.

4.6 4.4 1.3 ns

27.7 4.9 4.[ 24.9 30.5** 11.5'

25.6 26.9 5.5'

27.3 59.1 28.9 63.5 4.1 ns 9.4" 36.8 87.8 [9.4 37 468.7** 561.4"

5.5 795 0 0 499.7" 151.2" 2.9 475 1[15 6.9 25.9'· 81.2" 850 6.8 4.2 740 6.5' 2.3 ns

5.7 8 258.5" 6.1 7.6 109.1"

12.7 11.1 19.6" 11.1 12.6 16.9" 12.3 11.5 5.2'

5.2 4.8 39.0" 4.1 5.9 666.1" 2.6 37.6"

3

2.7 15.5"

3

1 0.7 370.0"

0.8 0.8 [ ns

16.0"

1.3

1.2 1.2 0.1 ns

6 6.7 27.7"

6.6 6.1 [6.9"

13.1 15.5 34.3"

15.1 13.5 12.6"

7.2 6.8 8.4" 5.1 8.9 861.6'-

Root

1.2 5.3 14.2 29.3 5 23.3 48.7 7.4 3 1.7 7 7.9 PS 10.8 6.4 4.7 2.7 0.4 0.7 5.6 14.3 23.3 4 32.8 76.1 6.2 DS 11.7 70.0" 48.4" 149.2'- 20.3-- 143.5*' 162.6" 0.1 ns 180.2** 54.7" 940.9" 595.3" 113.7" F-test 6.6' • Significant at p