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Influence of Drought Stress on Growth and Nodulation of Acacia origena

(Hunde) Inoculated with Indigenous Rhizobium Isolated from Saudi Arabia

Article · May 2015

DOI: 10.5829/idosi.aejaes.2015.15.5.12629

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American-Eurasian J. Agric. & Environ. Sci., 15 (5): 699-706, 2015
ISSN 1818-6769
© IDOSI Publications, 2015
DOI: 10.5829/idosi.aejaes.2015.15.5.12629

Influence of Drought Stress on Growth and Nodulation of Acacia origena (Hunde)

Inoculated with Indigenous Rhizobium Isolated from Saudi Arabia

Nader D. Shetta

Forestry and Wood Technology Department, Faculty of Agriculture, Alexandria University, Egypt

Abstract: Drought is the main abiotic factor affecting the survival of soil microorganisms and plant growth. The
present study was conducted at the Range and Forestry Applied Research Unit, King Saud University at Dirab,
South of Riyadh City to study the effect of drought stress on the growth and nodulation of Acacia origena
inoculated with indigenous Rhizobium. Three indigenous Rhizobium isolates (KS1, KS2 and KS3) were isolated
from the root nodules of seedlings of Acacia tortilis (Forssk.), Leucaena leucocephala (Lam.) and Acacia
saligna (Maslin), respectively. The tree seeds were obtained from Abha, Saudi Arabia and planted in a shade
house. Drought stress significantly reduced both seedling growth and nodulation of Acacia origena. Under
this stress, the Rhizobium isolates varied in their response to the drought period; they were able to create
nodules on the roots of the Acacia origena, while this declined with an increase in the drought period. A
treatment that involved withholding irrigation for 21 days was the most effective treatment for growth and
nodulation. Further, indigenous Rhizobium isolates KS1 and KS2 are drought tolerant. The inoculation of
seedlings with indigenous Rhizobium isolates might therefore improve the drought stress tolerance of Acacia
origena seedlings under these conditions.

Key words: Drought stress Indigenous Rhizobium Acacia origena Nodulation

INTRODUCTION of these species were described in a review of nine major

arid and semi-arid areas around the world. Studies of root
Drought is the main abiotic factor that affects the nodule bacteria under stress conditions in soil might
survival of soil microorganisms and plant growth and mimic their survival in natural habitats. This is because
drought stress has an adverse influence on water bacterial strains under carbon and energy limitations in
relations, mineral nutrition, metabolism and soil may be more sensitive to environmental stress than
photosynthesis [1]. Water is one of the most important they would be in rich laboratory media. In addition,
resources for agricultural production. In many parts of the rhizobia may withstand and successfully multiply under
world, water availability is economically and/or technically these conditions, but their infectibility and nodulating
limited; 28 and of the Earth’s land surface is considered to ability may be changed [14]. Selection of rhizobia that can
be too dry for crop production [2-4]. Drought and salinity withstand drought stress is important for improving plant
are considered by agronomic researchers to be challenges productivity for reforestation applications in desert and
for nodulating legume crops. However, there are many semi-desert areas. In Saudi Arabia, acacia populations
wild legumes that grow in deserts, some of which are play a major role in the productivity and stability of the
saline- and/or high pH-tolerant [5]. Studies on the desert environment, but they are threatened due to
nodulation of legumes have been conducted in many of indiscriminate cutting, extreme environmental conditions,
arid and semi-arid areas of Africa [6-10]. There are many such as drought and salinity and low density. There are
potential nodulated legumes native to arid and semi-arid much recent interest in the tolerance of wild legumes and
areas that have been studied for their tolerance to drought their symbionts to severe drought, salinity and elevated
or salinity, such as Prosopis spp. in the Americas, temperatures. However, natural symbiotic rhizobia with
Mimosa spp. in the Caatinga biome of Brazil, pasture legumes establish effective symbioses under these
species and trees, especially African acacias [11-13]. Some conditions [6]. Knowledge of the symbioses of woody

Corresponding Author: Nader D. Shetta, Forestry and Wood Technology Department,

Faculty of Agriculture, Alexandria University, Egypt. E-mail: n.shetta@yahoo.com.
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legume species in natural ecosystems and the diameter) containing a sterilized sand: vermiculate mix (2:1
distributions of root nodule bacteria under drought and v: v). The unstressed seedlings were watered every 3
salt stress conditions is limited. Because salinity is often days, while the drought-stressed seedlings were watered
associated with drought in semi-arid and arid regions, the every 7, 14 and 21 days. Each treatment had three
present study aims to examine the effect of drought stress replicates. Seedlings were inoculated with 10 ml of
on the growth and nodulation of Acacia origena Rhizobium isolates KS1, KS2, or KS3 containing
inoculated with indigenous rhizobia. approximately 1×109. Rhizobium cells grown in yeast
extract mannitol broth with shaking (200 rpm). Pots were
MATERIALS AND METHODS arranged in factorial experiment in complete randomized
design with three replicates in the greenhouse and the
Tree Species and Soil: Acacia origena are medium-sized experiment was conducted from October 2012 to the first
trees (6 meters height), native to Ethiopia, West Eritrea week of March 2013 and repeated from October 2013 to
and to regions across the Red Sea in Yemen and Saudi the first week of March 2014. Seedlings were maintained
Arabia. The bark is white to greyish-white in thick-papery at 28°C. At the end of the experiment, seedlings were
layers; younger branches have yellow to yellowish-brown harvested and analyzed for their height, diameter, nodule
bark. Leaves on reduced axillary shoots; pinnae usually 2- numbers, nodule dry weight and stem and root dry
4 pairs per leaf; leaflets with 14-34 pairs per pinna, weight.
glabrous, 6 mm × 2 mm. Inflorescence globular heads,
creamy-white, 1 cm in diameter, pedunculate, 2 cm long. Extraction and Analysis of Chlorophyll: Chlorophyll a, b
Pods are straight, purplish-green to red, 8 cm × 2 cm, and a+b were determined in fresh leaves by using the
obtuse or falcate, longitudinally and obliquely veined. method described by Porra et al. [16]. The leaf samples
The trees are restricted to only the eastern slopes of the were weighed and ground with 1 ml of solvent (N, N-
Asir and Abha regions and extend to an elevation of 2700 dimethylformamide (DMF)) using a pestle and mortar. The
meters together with Juniperus procera. The physical and homogenate, combined from a further three washings of
chemical characteristics of soil used in the study were as the pestle and mortar (each 1.5 ml) with the same solvent,
follows: the soil texture was sandy loam with pH 8.2 and was centrifuged at 2500 rpm in a bench centrifuge for 10
E.C 1.01 mmhos/m; total soluble cations (Na+, K+ and Ca++) min. The pellet was extracted with 1 ml of solvent in a
were 1.04,. 52 and 0.25 meq/l, respectively; organic matter homogenizer and the pooled supernatants were adjusted
1.0 and and total nitrogen 29.2 ppm. The soil was air-dried to a final volume of 8 ml. The spectrum was recorded
for 1 week, thoroughly mixed and autoclaved twice for 2 between 750 and 600 nm and the major red absorption
h (121 ~) on two consecutive days. peak was automatically determined using a UV-VIS
spectrophotometer (T80 UV/VIS Spectrophotometer, PG
Bacterial Strains and Isolation of Root-nodule Bacteria: instruments Ltd- USA) with the recording zeroed at 750
Rhizobium isolates KS1, KS2 and KS3 were isolated from nm. The Chl. a, b and Chl. a+b contents in µmol/l were
root nodules of seedlings of Acacia tortilis (Forssk.), calculated using the equations of Porra et al. [16].
Leucaena leucocephala (Lam.) and Acacia saligna
(Maslin), respectively, which were grown in the Dirab Statistical Analysis: The obtained data were submitted to
Valley, South of Riyadh City. The isolates were collected analysis of variance according to the method of Snedecor
from healthy, unbroken pink root nodules collected in the and Cochran [17] using ANOVA and LSD procedures
field. The method used to isolate root-nodulating bacteria available in the SAS software package (version 9.13,
from the seedlings was described by Vincent [15]. 2008).

Growth and Plant Material: The Acacia origena seeds RESULTS

used in this experiment were obtained from the Abha
region (South of the Kingdom) by the Range and Forestry Effect of Drought Stress on Growth Parameters and
Applied Research Unit, Faculty of Food and Agriculture Nodulation of Acacia origena: Drought is the main
Sciences, King Saud University. The seeds were immersed abiotic factor affecting the survival of soil microorganisms
in hot water (100°C) for 15 min and in cool water for 24 h, and plant growth. There were significant differences
then sown in plastic tray troughs containing a mixture of between the Rhizobium isolates and period of drought
sterilized vermiculate and sand (1:1 by volume). stress on seedling growth and nodulation of Acacia
Germination began at day 7. After four weeks, the origena over the two seasons. The differences in height,
seedlings were transplanted into plastic pots (15 cm diameter and number of nodules per plant between

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Fig. 1: The mean effect of Rhizobium isolates on height,

diameter and nodules number of Acacia origena
under drought period over two seasons.
Fig. 2: The mean effect of drought period on height,
Rhizobium isolates, drought stress and their interactions diameter and nodules number of Acacia origena
were highly significant at (P < 0.01) in both seasons, while over two seasons.
the other parameters varied between significant and not
significant in both seasons. The main effect of Rhizobium parameters. The seedlings in the 21 day drought period
isolates varied in their influence on seedling height and had the most impact compared with the control seedlings
diameter between KS1 and KS2 in the two seasons and in both growing seasons (Table 1). The seedlings
isolate KS3 produced the largest number of nodules (Fig. inoculated with the Rhizobium isolate KS3 were the most
1). The main effect of drought period followed the same tolerant to the drought treatments for height; the
trend as for Rhizobium isolates; the treatment irrigated percentage reduction for the two seasons was lower than
every 21 days was the most influential treatment with for KS1 and KS2; the reductions were 11% and 12%
regard to seedling height, diameter and the nodule after 7 days of withholding irrigation, respectively.
numbers (Fig. 2). After 14 days withholding irrigation, the height
The interaction between rhizobium isolates and reductions were 37% and 15%, while at 21 days the
drought period was significantly different for growth and height reductions were 52% and 15%, respectively
nodulation. Under drought stress, the isolates were able (Table 1). The same trend was found for nodule numbers;
to form nodules on the roots of the Acacia origena, but the Rhizobium isolate KS3 was the most tolerant to
this declined with an increase in drought period (Table 1). drought in both seasons. The percentage reductions in
Rhizobium isolates KS2 and KS3 were the most number of nodules per plant were 38%, 39% and 100% in
competitive under drought stress compared to KS1 in the the first season and 42%, 51% and 100% in the second
two seasons and showed improvement in some growth season for 7, 14 and 21 days of withholding irrigation

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Table 1: Interaction between Rhizobium isolates and drought period on the growth and nodulation of Acacia origena over two seasons
Seedling growth
---------------------------------------------------------------------------------------------------------------------------------------
First Season Second Season
Rhizobium Drought -------------------------------------------------------------------- ---------------------------------------------------------
isolates period (days) Height (cm) Diameter (mm) No Nod Height (cm) Diameter (mm) No Nod
KS1 Control 24.7 ±1.5a 1.62±0.37a 15.0±3.6a 31.0 ±3.6a 1.27±0.30b 11.0±1.0b
7 20.3±2.0b 1.57±0.05a 8.3±1.5bc 24.3±1.2b 1.20±0.19b 5.7±102d
14 20.0±2.6b 1.45±0.13a 7.0±1.0c 19.7±0.6bc 1.11±0.11b 3.0±1.0e
21 20.0±2.0b 1.44±0.16a 0.0±0.0d 17.7±1.5c 0.80±0.04c 0.3±0.2f
KS2 Control 20.0±2.0b 1.43±0.09a 10.3±1.5b 32.7±4.0a 1.57±0.15a 7.3±1.2cd
7 17.7±2.1bc 1.06±0.14b 7.0±1.0c 24.0±1.0b 1.53±0.08ab 6.7±1.5cd
14 14.7±1.6c 1.02±0.05b 7.3±1.5bc 19.0±2.6c 1.47±0.20ab 3.3±1.2de
21 13.0±0.9c 0.94±0.12bc 0.0±0.0d 17.0±4.3c 0.80±0.04c 1.0±0.5ef
KS3 Control 27.3±1.5a 1.30±0.13ab 16.0±3.0a 22.8±3.0bc 1.56±0.04a 14.3±4.0a
7 24.3±3.8a 1.29±0.12ab 10.0±2.6bc 20.0±2.6bc 1.47±0.20ab 8.3±1.5c
14 17.3±2.4ab 0.88±0.11bc 9.7±1.2bc 19.3±3.1bc 1.20±0.18b 7.0±1.0cd
21 13.0±0.9c 0.61±0.42c 0.0±0.0d 19.0±0.5c 1.06±0.07bc 0.0±0.0f
Means followed by the same superscript letter in each column are not significantly different at P<0.05 according to the LSD test. KS1, Rhizobium isolated
from Acacia tortilis seedlings; KS2, Rhizobium isolated from Leucaena leucocephala seedlings; and KS3, Rhizobium isolated from Acacia saligna seedlings.
No Nod: nodules number.

Table 2: Interaction effect between Rhizobium isolates and drought period on seedlings biomass of Acacia origena over two seasons
Seedlings biomass
------------------------------------------------------------------------------------------------------------------------------------------
First Season Second Season
Rhizobium Drought ------------------------------------------------------------------- ------------------------------------------------------------
isolates period (days) Nod Dry(mg) SD(g) RD(g) Nod Dry(mg) SD(g) RD(g)
KS1 Control 0.092±0.01a 0.24±0.01a 0.08±0.01a 0.034±0.010a 0.16±0.01a 0.063±0.010ab
7 0.023±0.004b 0.20±0.02b 0.07±0.01a 0.022±0.003b 0.10±0.07a 0.037±0.000c
14 0.022±0.007b 0.19±0.04bc 0.07±0.02a 0.016±0.005b 0.08±0.01a 0.030±0.005c
21 0.00±00c 0.17±0.01bc 0.07±0.02a 0.00±0.000c 0.07±0.01a 0.026±0.003c
KS2 Control 0.010±0.003c 0.07±0.02d 0.06±0.02b 0.094±0.012a 0.21±0.04a 0.083±0.006a
7 0.06±0.003c 0.03±0.0e 0.04±0.03cd 0.026±0.006b 0.20±0.02a 0.073±0.007abc
14 0.005±0.002c 0.02±0.01e 0.02±0.01e 0.024±0.022b 0.14±0.06a 0.070±0.022abc
21 0.00±00c 0.02±0.0e 0.03±0.01de 0.017±0.007b 0.11±0.05a 0.050±0.007abc
KS3 Control 0.023±0.005b 0.18±0.02b 0.07±0.01a 0.031±0.009b 0.19±0.04a 0.077±0.006a
7 0.018±0.009bc 0.17±0.03bc 0.06±0.02bc 0.026±0.006b 0.17±0.01a 0.060±0.000abc
14 0.013±0.002c 0.16±0.04c 0.05±0.00bc 0.022±0.007b 0.16±0.07a 0.053±0.012abc
21 0.00±00c 0.03±0.03e 0.03±0.01de 0.000±0.000c 0.12±0.09a 0.040±0.009b
Means followed by the same superscript letter in each column are not significantly different at P<0.05 according to the LSD test. KS1, Rhizobium isolated
from Acacia tortilis seedlings; KS2, Rhizobium isolated from Leucaena leucocephala seedlings; and KS3, Rhizobium isolated from Acacia saligna seedlings.
SD, Shoot dry weight (g); RD, Root dry weight (g); Nod D, nodular dry weight (mg).

compared with control seedlings, respectively. The seedling biomass. The nodule dry weight and
seedlings diameter varied between KS1 and KS2 shoot and root dry weight were reduced with
(Table 1). increasing drought treatments and 21 days of withholding
irrigation was the treatment with the greatest effect on
Effect of Drought on Seedlings Biomass of seedling biomass (Table 2). The Rhizobium isolates varied
Acacia origena: The interaction between Rhizobium in their tolerance to drought; the seedlings inoculated
isolates and drought stress periods was significant with rhizobium KS1 produced the highest number of
for shoot and root dry weight in both seasons. nodules, shoot and root dry weight in the first season,
Table 2 shows the mean values of the interaction while the Rhizobium isolate KSA2 had the highest number
between the rhizobium isolates and drought stress of nodules, shoot and root dry weight in the second
periods during the 1st and 2nd seasons with regard to season.

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Fig. 3: Mean effect of Rhizobium isolates on chlorophyll content over two seasons.

Fig. 4: Mean effect of drought period on chlorophyll content over two seasons.

Table 3: Interaction effect between Rhizobium isolates and drought period on chlorophyll content of Acacia origena over two seasons
Chlorophyll contents (µml/l)
------------------------------------------------------------------------------------------------------------------------------------------
First season Second Season
Rhizobium Drought -------------------------------------------------------------------- --------------------------------------------------------------
isolates period (Days) Chl. a Chl. b Chl. a+b Chl. a Chl. b Chl. a+b
KS1 Control 10.6±0.9a 3.47±0.29a 14.05±1.2a 10.3±1.13a 2.89±0.25ab 13.12±1.39a
7 8.36±0.46bc 2.71±0.11ab 11.06±0.55bc 9.65±1.28a 2.73±0.41ab 12.37±1.67ab
14 7.47±0.67c 2.07±0.09b 9.54±0.75c 4.65±0.80b 1.47±0.74bc 6.11±0.07c
21 2.30±1.0f 0.73±0.27c 2.03±1.2f 4.60±0.39b 1.18±0.09c 5.79±0.37c
KS3 Control 10.6±0.97a 3.47±0.29a 14.05±1.2a 7.95±4.47ab 3.10±0.44a 11.04±4.71ab
7 8.61±0.58b 2.88±0.17ab 11.46±0.73bc 7.94±4.27a 2.71±1.48ab 10.66±5.95ab
14 4.64±0.80d 1.47±0.74bc 6.11±0.06de 7.44±0.63bc 1.94±0.14bc 9.38±0.52bc
21 3.01±0.11ef 1.56±0.89b 4.57±0.9ef 2.64±0.65c 0.91±0.18c 3.55±0.83c
KS3 Control 9.72±1.68ab 2.73±0.46ab 12.45±2.1ab 10.23±1.14ab 2.89±0.26ab 13.12±1.39a
7 7.47±0.67c 2.07±0.09b 9.54±0.75c 9.63±01.31ab 2.70±0.45ab 12.32±1.75ab
14 4.46±0.70de 1.40±0.79bc 5.87±0.37e 7.49±0.70b 2.10±0.14b 9.59±0.83bc
21 2.08±0.30f 0.69±0.22c 2.77±0.52f 3.05±0.0.66c 0.99±0.12c 4.04±0.76c
Means followed by the same superscript letter in each column are not significantly different at P<0.05 according to the LSD test. KS1, Rhizobium
isolated from Acacia tortillis seedlings; KS2, Rhizobium isolated from Leucaena leucocephala seedlings; and KS3, Rhizobium isolated from
Acacia saligna seedlings.

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Effect of Drought Stress on Chlorophyll Content of tortilis and Acacia gerrardii subsp negevensis and
Acacia origena: The chlorophyll content is one of the Ramos et al. [23] on common bean. The mechanisms for
most significant parameters related to the physiological conserving water adopted by acacia trees to endure
status of plants and estimations of chlorophyll content water stress in this study seem to be related to reducing
and related chlorophyll parameters can be used as indexes absorption via thin roots and decreased leaf area. The
of nutrient status, physiological stress and changes in results show that under different drought stress
abiotic factors. The analysis of variance for Acacia periods, stem diameter was reduced with increasing the
origena detected variation between Rhizobium isolates, period of withholding irrigation compared with normal
drought period and their interaction in both seasons. irrigation. This is in agreement with those obtained by
Chlorophyll a and a+b were significantly different, Aref and El- Juhany [21, 22]. The decreased in root dry
whereas chlorophyll b was not significantly different in weight under drought stress conditions presented in our
the first season, while in the second season, the drought study concurs with other results for woody species in the
period differed significantly. The mean effect of seedling stage, such as reported by Ibrahim [24] on
chlorophyll content in the seedlings inoculated with poplar, Aref and El-Juhany [22], on some acacia species
Rhizobium isolate KS1 was high compared with KS2 and and El-Juhany and Aref [19] on Leucaena spp. However,
KS3 in both seasons (Fig. 3). The drought period had a the lower percentage of reduction in shoot and root dry
significant effect on the chlorophyll content of the weight may reflect the morphological plasticity of root
seedlings. The chlorophyll content was reduced with systems that enable them to cope with variable soil
increasing irrigation intervals (21 days) (Fig. 4). The conditions.
interaction between Rhizobium isolates and drought The reduction in chlorophyll content during seedling
stress periods for chlorophyll a and a+b was significant exposed to different drought stress periods may be
in the first season, while in the second season was not attributed to the decrease in number of leaves. The
significant. Rhizobium isolate KS2 was most tolerant to reduction of chlorophyll content related to the reduction
drought stress, followed by KS1 and KS3 was most of leaf area as a result of drought stress. This result is in
sensitive to drought in the 1st and 2nd growing seasons agreement with the results obtained by Kozlowski et al.
(Table 3). The reduction percentages in the chlorophyll [25], who indicated that water deficits reduced leaf area by
content of seedlings inoculated with KS2 in the first inhibiting the initiation of leaves, as well as their
season were 18.8%, 17% and 18.4% for chlorophyll a, b subsequent enlargement. Also, Aref and El-Juhany [22]
and a+b, respectively. The same trend was found in reported that water stress reduced leaf area, as well as
the second season; the reduction in chlorophyll a, b reduced the surface conducts photosynthesis, thus
and a+b was lower than the in first season, the decreasing the quantity of photosynthate available for
percentage reductions were 1%, 13% and 3.4%, plant growth. In non-irrigated soil, the seedlings were not
respectively (Table 3). able to produce a new root or root tips because the water
deficiency lowered the growth activity of the seedling
DISCUSSION AND CONCLUSION after the nursery phase. Additionally, the root nodules
become thickened, larger and more resistant to infection
The reduction in the growth of the seedlings due to with Rhizobium [26]. Furthermore, results indicated that
drought encompassed all growth characteristics Rhizobium isolates varied in their response to drought
measured in the present study. Many aspects of plant period, but the treatment of withholding irrigation for 21
growth are sensitive to drought stress in both cellular days had the most impact on nodulation. Additionally, it
and whole-plant [18]. Decreased stem height and was found that the indigenous rhizobium isolates can be
diameter of woody trees under drought stress conditions more tolerant to drought stress. The most tolerant
was reported also by El-Juhany and Aref [19]. The rhizobium isolates were KS1 and KS2. This result was in
results of this study indicated that drought period contrast with the findings of Swaine et al. [27], who
significantly reduced the seedling height and shoot and reported that the symbiosis of legume-Rhizobium, in
root dry weight in Acacia origena. A decrease in shoot general, is known to be more sensitive to environmental
dry weight due to drought stress was inevitable such as stress (especially drought) than the uninfected legume
the reduction in both height and diameter. Similar Rhizobium. Drought tolerant strains of the bacterium have
observations were reported by Pokhiryal et al. [20] on been documented for both crop species and agroforestry
Acacia nilotica, Aref and El Guhany [21, 22] on Acacia tree species by several studies. Nodulation and nitrogen

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 Am-Euras. J. Agric. & Environ. Sci., 15 (5): 694-698, 2015

fixation in legume-Rhizobium associations are adversely 2. Bray, E.A., 2004. Genes commonly regulated by
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8. Wolde-meskel, E., T. Berg, N.K. Peters and
nodules on the seedlings of Acacia origena under
A. Frostegård, 2004. Nodulation status of native
drought stress. Thus, indigenous Rhizobium strains can
woody legumes and phenotypic characteristics of
help the Acacia tress to withstand the stress conditions
associated rhizobia in soils of Southern Ethiopia.
found in the field. Our results suggest that the selection
Biology and Fertility of Soils, 40: 55-66.
of Rhizobium isolates with tolerance to drought stress is
9. Odee, D.W., J.M. Sutherland, E.T. Makatiani,
essential for reforestation and improving the productivity
S.G. McInroy and J.I. Sprent, 1997. Phenotypic
of Acacia trees under the natural conditions of Saudi
characterization and composition of rhizobia
Arabia. Even if these abiotic factors do not act alone in
associated with woody legumes growing in diverse
natural conditions, it is necessary to test the efficiency of
Kenyan conditions. Plant Soil, 188: 65-75.
the Rhizobium isolates under shade house conditions 10. Nick, G., P. de Lajudie, B.D. Eardly, S. Soumalainen,
prior to using them as inoculants in the field. L. Paulin, X. Zhang, M. Gillis and K. Lindström,
1999. Sinorhizobiu marboris sp. nov and
ACKNOWLEDGMENTS Sinorhizobium kostiense sp. nov., isolated from
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