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This article was downloaded by: [York University Libraries]On: 13 November 2014, At: 09:52Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office:Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK
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Effects of Simulated Acidic Precipitation andAluminum on the Growth of Alnus glutinosaand Alnus hirsuta Seedlings and Their NitrogenFixationD. K. Lee a , C. H. Yun b , M. I. Choi b & Haworth Continuing Features Submissiona School of Forest Sciences & Forest Products, Seoul, 151-742, Koreab Graduate Research Assistant, Department of Forest Resources, Seoul NationalUniversity, Seoul, KoreaPublished online: 25 Oct 2008.
To cite this article: D. K. Lee , C. H. Yun , M. I. Choi & Haworth Continuing Features Submission (1994) Effectsof Simulated Acidic Precipitation and Aluminum on the Growth of Alnus glutinosa and Alnus hirsuta Seedlings andTheir Nitrogen Fixation, Journal of Sustainable Forestry, 1:3, 71-92, DOI: 10.1300/J091v01n03_06
To link to this article: http://dx.doi.org/10.1300/J091v01n03_06
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Effects of Simulated Acidic Precipitation and Aluminum on the Growth of Alnus glutinosa and Alnus hirsuta Seedlings
and Their Nitrogen Fixation
D..K. Lee C. H. Yun M. I. Choi
ABSTRACT. This study was conducted to examine the effect of simulated acidic precipitation and aluminum treatment on the growth of Alnus species and the contents of nitrogen fixed by Fran- kia. The growth and nitrogen contents of Alnus species inoculated with nitrogen-fixing organisms were greater than those of non- inoculated Alnus species when treated with simulated acidic preci- pitation. The nitrogen contents were higher in the soils used for the growth of Alnus glurinosa inoculated with Frankia than in those without Frankia inoculation when treated with either acidic preci- pitation or aluminum. The-development of root hairs treated with simulated acidic precipitation was poor as lhe pH level decreased, and the injury of A. glurinosa was more severe than that of Alnus hirsuta under scanning electron microscopy (SEM). The growth and nitrogen contents of A. hirsura inoculated with Frankia were greater than those of non-inoculated species and A. glutinosa when treated with aluminum. The leaves of A. glutinosa became yellowish-brown and fell earlier. Under SEM and light microscopy, the surface layer of roots in both Alnus species was injured severely and the number of root hairs decreased as aluminum levels increased.
D. K. Lee is Professor, Department of Forest Resources, Seoul National Uni- versity. Suwon 441-744, Korea C. H. Yun and M. I. Choi are Graduate Research Assistants at the Department of Forest Resources, Seoul National University. '
Journal of Sustainable Forestry, Vol. l(3) 1993 O 1993 by The Haworth Press, Inc. All rights reserved. 71
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JOURNAL OF SUSTAINABLE FORESTRY
INTRODUCTION
Actinorhizal plants such as Alnus, Elaegnus, Myrica, Casuarina and others are ecologically and/or economically important to refor- estation and land reclamation programs due to nitrogen fixation associated with the genus Frankia (Fessenden 1979, Huss-Danell 1986). Their contributions to wood and fiber production have been seen to be very large (Miller and Murray 1979, Zavitkovski et at. 1979). Such examples are often found at hillsides in Korea.
The acidity of precipitation has increased in various regions throughout the world during the last two decades (Bolin et al. 1971, Cogbill and Likens 1974, Lee and Kim 1990, Liens and Bormann 1974). Thus, the effects of acidic precipitation derived mostly from sulfur dioxide and oxides of nitrogen on soils and vegetation are numerous and complex: an increase in leaching of nutrients from plant leaves and soils; an inhibition or stimulation of certain plant diseases; foliar injury; effects on seed germination, seedling emer- gence and the growth of plants; and an inhibition of the nitrogen- fixing activities in legumes (Dochinger and Seliga 1976, Evans et al. 1977, Jonsson and Sundberg 1972, Lee and Kim 1986, Lee and Weber 1979, Reuss 1975, 1978, Weber and Lee 1977, Wood and Bonnann 1975). However, sulfur added in the fertilizer or manure plus H2S04 resulted in an increased yield for certain crops (Cates et al. 1984, Jones et al. 1979).
In a forest, the long-term impact of acidic precipitation may result in soil acidification, aluminum mobilization and aluminum toxicity. Plant growth or particularly the growth of root length was damaged by low concentration of aluminum (Alva et al. 1986, Foy 1988). There are a few known aluminum injuries: (1) an inhibition of cell division and growth in roots by hindering the uptake of water and nutrients; (2) a collapse of plant tissues due to the interaction between aluminum and essential microelements such as Ca, Mg, or P; and (3) respiration damages by replacing toxic substances of aluminum into respiring organs.
Korea is also experiencing diverse pollutions in air, water and soils (Lee and Kim 1990). To take measures against such damages for the future, various basic and applied research on the impacts from pollutants is needed.
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Lee, Yun, and Choi 73
Thus, the objectives of this study were to examine the effects of simulated acidic precipitation and aluminum treatment on the growth of Alnus glutinosa and Alnus hirsuta seedlings and their nitrogen fixation.
MATERIALS AND METHODS
Seed Collection, Seedling Growth and Frankia Inoculation
Seeds were collected in October 1990 from 9-year-old-trees of Alnus hirsuta Rupr. growing at the Forest Nursery of Seoul National University, and from 10-year-old trees of Alnus glurinosa (L.) Gaertn. growing at the Institute of Forest Genetics, Suwon, Korea.
The seeds were sown in a plastic container containing sand after autoclaving at 10S°C, and the seedlings were grown for two months. Then each of the seedlings were transplanted into a pot (500 cm3) containing sand and grown for one month.
Based on daily growth rate, complete nutrients (Tables 1 and 2) reported by Ingestad and Lund (1979) were given daily to the seedlings during the experimental period, but nutrients without only nitrogen given to those inoculated with Frankia.
~~e s&lings of A. glutinosa grown two weeks after germination were inoculated with either Frankia AvcIl or crushed nodule meth- od (Lim and Lee 1989). However, those of A. hirsuta were inocu- lated only with crushed nodule method. Nodules for inoculation were collected from the roots of parent trees from which the seeds were collected. Three-month-old seedlings of two Alnus species were used for the studies of acidic precipitation and aluminum treatments. All of the experiments were conducted in a greenhouse.
Simulated Acidic Precipitation and Aluminum Treatment
Simulated acidic solution was prepared by a mixture of sulfuric and nitric acids (H2S04 : HN03 = 3:1, vlv). The sulfuric and nitric acids were diluted with tap water and adjusted to pH 3.0, 4.0, 5.6 and 6.4. Tap water (pH 6.4) was treated as control.
Forty milliliters of acidic solution were treated twice per week to
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74 JOURNAL OF SUSTAINABLE FORESTRY
TABLE 1. Composition of nutrient solutions used for the growth of Ainus species
Stock solution A (glL) Stock solution B (g/L)
Material Concentration Material Concentration
KSOc 24.48 Ca(N0s)z. 4H7.0 20.66 WzWz 33.48 Mg(No3)z. 6H7.0 44.88 K W 4 7.75 HNQ(1N) 20ml KNOJ 15.32 Micro 8406+ 25101 WJ 113.67
+ Micro 8406 solution
Material Concentration (g/O. 5 L)
Fe(N4)3. 9 M 50.64 b(N03)z. 4MO 18.28 Zn(NQ)z. 4 M 2.399 CuC12.2HzO 0.805 NaW4.2HzO 0.1765 NazB& 9.307 WOd65.6b) 10.55
each of the uninoculated plants from 6 August to 25 October 1991. During this period, the plants were harvested biweekly. Then, the height, root collar diameter and leaf number of the plants were measured. The contents of total nitrogens in both plants and soils were also measured after drying in an oven at 80" C.
A combination of aluminum chloride and aluminum sulfate (AlCl3 .4H20 : A12(S02)3 . 18H20 = 1: 1, v/v) was used to conduct the experiment of aluminum (Al) treatment (McConnick and Steiner 1978). A 0.004M calcium nitrate (Ca(N03)2.4H20) was included in the treatment solutions. Then, these two solutions were added to the nutrient solution described in Table 1 and 2. Forty milliliters of all the solutions were treated weekly to each of the seedlings at A1 concentrations of 0,50,100 and 200 mgb .
The techniques for seed sowing and growth of the seedlings inoculated with Frankia Avcn for the experiment of acidic preci-
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Lee, Yun, and Choi 75
TABLE 2. Contents of the elements contained in nutrient solutions
Element Contents(%)
pitation and aluminum treatments were the same as those used in uninoculated seedlings.
The seedlings of A. glutinosa inoculated with Frankia Avcn were treated separately with simulated acidic precipitation or aluminum starting 27 January 1992 for two months, and those of both species of Alnus inoculated with crirshed nodule method were treated sepa- rately with simulated acidic precipitation or aluminum starting from 6 February 1992 for two months. During this period, the plants were harvested biweekly. All the measurements were the same as those used for uninoculated plants.
RESULTS AND DISCUSSION
Effects of Simukkted Acidic Precipitation on Seedling Growth and Nitrogen Fixation
Growth of height and root collar diameter and number of leaves in A. glutinosa seedlings treated with simulated acidic precipitation are shown in Figures 1,2 and 3.
A. glutinosa seedlings showed better growth of height and root collar diameter at pH 5.6 and pH 6.4 than at other pH levels whether those were inoculated with Frankia or not. However, the growth of
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76 JOURNAL OF SUSTA1NABL.E FORESTRY
FIGURE 1. Effects of simulated acidic precipitation on the height growth of Alnus glutinosa and Alnus hirsuta seedlings ( A, 6: no inoculation, C, D: inoculated with crushed nodule, E: inoculated with Frankia Avcll).
140
e a 120
110
loo 1st 2nd 3rd 4th 5th Harvest time
Alnus glutinosa
Harvest time Alnus hirsuta
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Lee, Yun; and Choi 77
FIGURE 2. Effects of simulated acidic precipitation on the root collar diame- ter growth of Alnusglutinosa and Alnus hirsuta seedlinas ( A. B: no inocula- tion, C, D: inoculated with crushed nodule, ~:inoculated;vilh kankia~vcl l ) .
Harvest time AInus hirsuta
Harvest time Alnus glutirmsa
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78 JOURNAL OF SUSTAINABLE FORESTRY
FIGURE 3. Effects of simulated acidic precipitation on the leaf number of Alnus glufinosa and Alnus hirsuta seedlings ( A, B: no inoculation, C, D: inoculaled with crushed nodule, E: inoculated with Fmnkia Avcll).
c
15
c
Ha~est time A l m s hirsute
- p n 3 . 0 4 - p H 4 . 0
+-pH 5.6 -c- 6.4
Hawest time
Alnus glutinosa
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Lee. Yun, and Choi 79
uninoculated seedlings as well as nitrogen contents in both plants and soils (Tables 3 and 4) was good at pH 4.0. This could mean that acidic precipitation including nitric acid might stimulate plant growth by the action of nutrient supply (Lee and Weber 1979, Ogner 1980). The nitrogen contents of A. glurinosa seedlings treated at pH 4.0 were greater than those treated at other levels of pH (Table 3). Those treated at pH 3.0 exhibited the poorest growth, which indicates that the seedlings might be damaged by acid treatment.
The height growth of A. glutinosa seedlings inoculated with crushed nodules decreased as pH levels decreased (Figure 1-C).' Trends similar to height growth were shown in nitrogen contents of plant tissues. Thus, it was considered that height growth was retarded at early stages as acidity increased.
The seedlings inoculated with either Frankia AvcIl or crushed nodules showed better height growth than uninoculated ones. It was apparent that the seedlings inoculated with Frankia AvcIl grew well even at low pH levels. However, root collar diameter growth and nitrogen contents of the seedlings inoculated with crushed nod- ules were rather greater than those inoculated with Frankia Avcn (Table 3).
The number of leaves of the uninoculated seedlings decreased as
TABLE 3. Nitrogen contents (% of the total dry weight per seedling).of the Alnus species grown under the various pH treatments
pH levels A. glutinose A. hirsute
without 6.4 -8
Frankia 5.6 0.618 0.905 inoculatian 4.0 0.863 0.785
3.0 0.513 0.768
inoculated 6.4 1.370 1.375 with crvshad 5.6 1.205 1.343 nodules 4.0 0.833 0.718
3.0 0.820 1.018
inoculated 6.4 1.212 with Frankia 5.6 1.234 AvcIl 4.0 1.046
3.0 1.008
8 No data available
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80 JOURNAL OF SUSTAINABLE FORESTRY
TABLE 4. Nitrogen contents (%)of the soils under the various pH treatments
pH levels A. glutinma A. hirsuta
without 6.4 -E
Frankia 5.6 1.065 0.713 imculation 4.0 1.423 0.880
3.0 0.833 0.730
inoculated 6.4 I. 120 0.775 with crushed 5.6 0.863 0.819 nodules 4.0 0.907 0.800
3.0 0.888 0.798
inoculated 6.4 1.498 with Frenkia 5.6 1.278 Avcil 4.0 1.232
3.0 1.060
8 No data available
acidity increased, and leaves fell earlier at the lowest pH level during the period of experiment. The result that the leaves treated with acidic precipitation fell earlier is similar to that reported by Jacobson et al. (1980) who found early leaf-falls and yield reduc- tion in various crop plants treated with acid rain.
The nitrogen contents of the soils for the growth of uninoculated Alnus seedlings decreased as the levels of pH decreased, and were also lower than those inoculated with Frankia (Table 4). This in- crease in nitrogen contents of the soils where inoculated seedlings grew might be explained due to exudation of nitrogen fixed by Frankia to the soils from the roots.
At early stages, the height growth of A. hirsuta seedlings without Frankia inoculation was rather better at low levels of pH than at higher levels (Figure 1-B), which may be explained by the uptake of nitrogen itself contained in acidic precipitation during the early growing period (Cates et al. 1984, Cole and Johnson 1977). After that, the growth was retarded due to greater acidity in soils.
The early growth of root collar diameter in A. hirsuta seedlings without Frankia was the best at pH 4.0 (Figure 243). The number of leaves and nitrogen contents of A. hirsuta seedlings without Fran- kia decreased as acidity increased (Figure 3-B, Table 4). However,
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Lee, Yun. and Choi 81
the A. hirsuta seedlings inoculated with crushed nodules showed poor height growth, root collar diameter, leaf number and nitrogen contents of the plants at low pH levels. The nitrogen contents of the soils where A. hirsuta seedlings were grown showed no differences between with and without Frankia inoculation, but those of the seedlings inoculated with Frankia were greater than those of unin- oculated ones. This probably resulted from nitrogen fnation.
Effects of Aluminum on Seedling Growth and Nitrogen Fixation
The growth of height and root collar diameter, and number of leaves in A. glutinosa seedlings treated with various concentrations of aluminum are shown in Figures 4, 5 and 6. The growth and
. nitrogen contents of uninoculated A. glutinosa seedlings decreased as the concentration of aluminum increased. In particular, there were no differences in the nitrogen content of the seedlings between the two concentrations of 0 and 50mgL. but significant decrease was shown at the concentrations of 100 and 200 m g n (Table 5). Although aluminum toxicity is usually exhibited in roots of the plants, no significant damages are shown at low aluminum concentration in this experiment. 'Ihese results are similar to that reported by Foy (1988) who found there were no aluminum toxicities at low concentrations.
The growth of the seedlings inoculated with either crushed nod- ules or Frankia Avcn showed rather better growth at low concentra- tions (Figures 4-C, D, E and 5-C, D, E). However, poor growth was shown at high concentrations. Inoculation seedlings of both species were better in growth of height and root collar diameter, and nitro- gen content of the seedlings.
The number of leaves in the seedlings of both species decreased as the aluminum concentrations increased. Greater reductions were found at high concentrations, which resulted from leaf-drying and leaf-falling due to aluminum effects (Figure 6). Such great reduc- tions in leaf numbers were apparent in the uninoculated seedlings. The leaves of those seedlings became whitish yellow when continu- ing to be treated with 200rngfL.
Less leaves were lost of inoculated seedlings than of uninoculat- ed ones, which can be explained by increased tolerance to alumi- num due to nitrogen fixed by Frankia. Aluminum toxicity became
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82 JOURNAL OF SUSTAINABLE FORESTRY
FIGURE 4. Effects of aluminum on the height growth of Alnusglutinosa and Alnus hlrsuta seedlings (A, 8: no inoculation, C, D: inoculated with crushed nodule, E: inoculated with Frankia Avcll).
ti la,
110
1st 2nd 3rd 4th 5th 180
1 i'O
160 E 0 150 -
140
5, 130
b 120
110
1st 2nd 3rd 4th 5th 6th
Harvest tlme
AInus hirsuta
Harvest tlme
Alnus glutinosa
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Lee, Yun, and Choi 83
FIGURE 5. Effects of aluminum on the root collar diameter growth of Alnus glutinosaand Alnus hirsutaseedlings (A. B: no inoculation. C. D: inoculated with crushed nodule, E: inoculatedwiih Frankia Avcll).
Harvest tlme
Alr~us hirsuta
Harvest time
Alnus glutinosa
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84 JOURNAL OF SUSTAlNABLE FORESTRY
FIGURE 6. Effects of aluminum on the leaf number of Alnusglutinosa and Alnus hirsuta seedlings ( A, 8: no inoculation, C, D: inoculated with crushed nodule, E: inoculated with Frankia Avcll).
Harvest time Alnus hirsuta
-0- 100 WlL -0- 200 ng/L
0J 11;t I
3rd 5th Harvest time
Alnus glutinosa
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Lee, Yun, and Choi 85
TABLE 5. Nitrogen contents (% of the total dry weight per seedling) of the Ainus species grown under the various aluminum levels
Aluminium levels A. glutinosa A. hirsute
without 0 mg/L 0.756 0.982 Frankia 50 mg/L 0.776 0.816 inoculation 100 mg/L 0.634 0.708
200 mg/L 0.498 0.564
inoculated 0 mg/L 1.300 1.170 with crushed 50 mg/L 1.308 1.123 nodules 100 mg/L 1.078 0.950
200 mg/L 0.948 0.968
inoculated 0 mg/L I. 402 -P
with Frankia 50 mg/L 1.300 - AvcIl 100 mg/L 1.092 -
200 mg/L 0.950
* No data available
severe when soil acidity was increased, which may stimulate early leaf falls or growth damages (McFee 1983, Poinke and Corey 1967). The nitrogen contents of the plants were usually reduced at high concentrations of aluminum.
The growth of height and root collar diameter in uninoculated A. hirsuta seedlings was good at various concentrations of aluminum except for the concentration of 200mgL (Figures 4-B and 5-B).
The leaves of A. hirsuta seedlings fell much at high concentration of aluminum, and such leaf-falls were true of uninoculated seedlings.
There were no differences in the height and root collar diameter growth and leaf number of A. hirsuta seedlings inoculated with crushed nodules among various aluminum concentrations (Figures 4-D, 5-D and 6-D). In particular. tolerance to aluminum was exhib- ited even at the concentration of 200mgIL. However, inoculated seedlings of A. hirsuta when treated with 200mgiL of aluminum seemed to absorb more nitrogen from the s o h than those of A. glutinosa as less nitrogen contents were formed in the soils for the growth of A. hirsuta (Table 6). Furthermore, the nitrogen contents
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TABLE 6. Nitrogen contents (%)of the soils by the various aluminum levels
Aluminium levels A. glutinose A. hirsute
without 0 mglL 1.072 1.226 Frankie 50 mg/L 0.850 0.972 inoculation 100 mglL 0.758 0,780
200 mg/L 0.662 0.650 - - --
inoculated 0 mg/L 0.928 1.030 with crushed 50 mglL 0.938 0.915 nodules 100 mg/L 0.943 0.865
200 mg/L 0.825 0.713
inoculated 0 mg/L 1.076 -4 with Frankis 50 mglL 0.980 - AvcIl 100 mglL 0.704 -
200 mg/L 0.654
* No data available
of Alnus seedlings inoculated with crushed nodules remained constant from low to high concentrations of aluminum. Thus, A. hirsuta seedlings when inoculated with Frankia showed greater tolerance to aluminum than A. glutinosa seedlings on the basis of their growth performance and nitrogen uptake.
The uninoculated seedlings of A. hirsuta were less tolerant to aluminum, but its inoculated seedlings were very tolerant without leaf falls. There were less vatiations in nitrogen contents of the seedlings inoculated with Frankia, whereas there was much varia- tion in uninoculated seedlings.
Root Damage Due to Simulated Acidic Precipitation and Aluminum
The root hairs and epidermal tissues of A. glutinosa seedlings inoculated with Frankia were developed well at pH 5.6 and pH 4.0 (Figures 7-B and -D), but they were poor with damage in epidermal tissues at pH 3.0 (Figure 7-F). Those of A. hirsuta seedlings inocu- lated with Frankia were similar to the result of A. glutinosa at pH 4.0. At pH 3.0 of simulated acidic precipitation, its damage was
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FIGURE 7. Light microscopy of the roots in Alnusseedlings treated with acidic precipitation. (A) A. hirsuta roots inoculated with Frankia aftertreatment with pH 5.6 for 4 weeks. (B) A. glutinosa roots inoculated with Frankiaafter treatment with pH 5.6 for4 weeks, (C) A. hirsuta roots inoculated with Frankia after treatment with pH 4.0 for 4 weeks, (0) A. glutinosa roots inoculated with Frankia after treatment with pH 4.0 for 4 weeks, (E) A. hirsuta roots inoculated with Frankiaafter treatment with pH 3.0 for 4 weeks, (F) A. glutinosa roots inoculated with Frankia after treatment with pH 3.0 for 4 weeks. (H: root hair, T: epidermal tissues, DH: damaged root hair. DT damaged epidermal tissues, LM X 200).
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Lee, Yun, and Choi 89
observed but was less than that in A. glutinosa roots (Figure 7-E), which may be explained by more hairs and better arrangement of root in A. hirsuta-roots. The roots of inoculated A. glutiiosa seed- lings without aluminum treatment were bar-shaped, healthy and curled in part (Figures 8- and -C). However, they were rough at the surface layer of root hairs when treated with SOmgIL, and the dam- age became severe as the concentration of aluminum increased (Figures 9-B and -D). At the concentration of 200mg.L only a few root hairs were formed and those were much damaged at epidermal tissues in both species. However, A. glutinosa showed more severe damage than A. hirsuta. Such damages were similar to those reported by McQuattie and Schier (1990). and Schier (1985).
Therefore, the treatments with acidic precipitation and aluminum did damage to both hairs and epidermal tissues of the roots. How- ever, the damage due to aluminum was greaterthan that due to acidic precipitation.
CONCLUSIONS
The effect of simulated acidic precipitation and aluminum on the growth of Alnus species and the nitrogen contents fixed by Frankia was examined.
The growth and nitrogen content of Alnus species inoculated with Frankia was greater than those of uninoculated Alnus species when treated with acidic precipitation. The nitrogen content was higher in the soils used for the growth of A. glutinosa inoculated with Frankia than in those without Frankia inoculation when treated with either acidic precipitation or aluminum. The develop- ment of root hairs treated with acidic precipitation became poor as the pH level decreased, and the injury of A. glutinosa was more severe than that of A. hirsuta under SEM.
The growth and nitrogen contents of A. hirsuta inoculated with Frankia were greater than those of uninoculated species and A. glutinosa when treated with aluminum. The leaves of A. glutinosa became yellowish-brown and fell earlier.
Under the SEM and light microscopy, the surface layer of roots of both Alnus species was injured severely and the number of root hairs decreased as aluminum levels increased.
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FIGURE 9. h h t microscopy of the roots jn Ahusseedlings treated with different aluminum concentrations. (A) A. hirsuQ mots inoculated with Frankia after treatment wth 100mgIL for 4 weeks, (B) A. glutinosa roots inoculated with Frankia after treatment with 100 mg/L for 4 weeks, (C) A. hrisuta mots inoculated with Fmkia after treatment with 20OmgJL for 4 weeks, (D) A. glutimsa roots inoculated with Frankia after treatment with 200mglL for 4 weeks. (H: root hair, T epidermal tissues, DH: damaged root hair, DT damaged epider- mal tissues, LM X 200).
-- - A - -- A H .p- H
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Lee, Yun, and Choi
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