全 文 ：丛枝菌根真菌对百喜草的生理特性的影响*
肖家欣1,2, 任摇 群1, 吴雪俊1, 陈迎迎1, 张绍铃2
(1 安徽师范大学生命科学学院, 生物环境与生态安全安徽省高校省级重点实验室, 安徽 芜湖摇 241000;
2 南京农业大学园艺学院, 江苏 南京摇 210095)
摘要: 采用盆栽法研究了丛枝菌根 (AM) 真菌摩西球囊霉 (Glomus mosseae) 对水分胁迫条件下百喜草
(Paspalum notatum) 生长、 渗透调节及抗氧化酶的影响。 结果表明: 接种 AM真菌显著提高了百喜草的株
高、 地上部与根部鲜重、 地上部 P、 K、 Mn及根部 P、 Ca、 Mn含量, 明显降低了地上部 Zn及根部 Fe、 B、
Cu水平; 随着干旱程度的加深, 接种株的地上部相对含水量及叶绿素含量相对稳定且均显著高于未接种
株, 接种株地上部相对电导率、 MDA含量均显著低于未接种株, 接种株的地上部 POD 活性与脯氨酸含量
均显著增加且均显著高于未接种株, AM侵染对 SOD活性的影响较小。 可见, 接种 AM真菌 Glomus mosse鄄
ae提高了植株体内保护酶活性 (如 POD) 及渗透调节能力 (如脯氨酸、 P、 K、 Ca等渗透调节物含量的增
加), 从而显著增强了百喜草的抗旱性。
关键词: 丛枝菌根真菌; 百喜草; 抗氧化物质; 渗透调节; 水分胁迫
中图分类号: Q 945摇 摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2011)05-521-08
Effects of Arbuscular Mycorrhizal Fungi on Physiological Character
of Bahia Grass (Paspalum notatum, Poaceae)
XIAO Jia鄄Xin1,2, REN Qun1, WU Xue鄄Jun1, CHEN Ying鄄Ying1, ZHANG Shao鄄Ling2
(1 Key Laboratory of Biotic Environment and Ecological Security, Anhui Province, College of Life Sciences,
Anhui Normal University, Wuhu 241000, China; 2 College of Horticulture,
Nanjing Agricultural University, Nanjing 210095, China)
Abstract: The effects of arbuscular mycorrhizal (AM) fungus, Glomus mosseae, on growth, osmotic adjustment
and antioxidant enzymes of bahia grass (Paspalum notatum) were studied in potted plants under water stress condi鄄
tions. AM colonization significantly enhanced the plant height, root and shoot fresh weight, Phosphorus (P), pota鄄
sium (K), manganese (Mn) contents in shoots, P, calcium (Ca), Mn contents in roots, whereas obviously de鄄
creased zinc (Zn) content in shoots, iron (Fe), boron (B), copper (Cu) contents in roots. During water stress,
the relative water and chlorophyll contents were relatively stable and signifciantly higher in AM than in non鄄AM
plants, AM inoculation notabley decreased the shoot relative conductivity and malondialdehyde (MDA) content,
markedly increased shoot peroxidase (POD) activity and proline content, while AM infection did not affect the dis鄄
mutase (SOD) activity of shoots. Our results suggested that AM colonization improved the protective enzyme activity
(such as POD) and osmotic adjustment originating from proline P, K, Ca, resulting in the enhancement of drought
tolerance.
Key words: Arbuscular mycorrhizal fungi; Bahia grass; Antioxidant; Osmotic adjustment; Water stress
植 物 分 类 与 资 源 学 报摇 2011, 33 (5): 521 ~ 528
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 3724 / SP. J. 1143. 2011. 11007
* Foundation items: Specialized Research Fund for the Doctoral Program of Higher Education (20103424120002), China Postdoctoral Sci鄄
ence Foundation (20080430172) and Anhui Provincial Natural Science Foundation (070411004)
Received date: 2011-01-10, Accepted date: 2011-08-02
作者简介: 肖家欣 (1970-) 男, 博士, 副教授, 主要从事植物生理与生态方面的研究。 E鄄mail: xjx0930@ 163. com
摇 Drought stress limits production of crop plants,
particularly in arid and semi鄄arid areas ( Kramer
and Boyer, 1997). Arbuscular mycorrhizal (AM)
fungal symbiosis is a mutualistic association between
AM fungi and the roots of terrestrial plants, the
ancient fungi can colonize approximately 80% of the
Earth爷s land plant species, it爷 s well documented
that adaptive AM fungi inoculation can enhance the
drought tolerance of host plant such as citrus (Wu
et al., 2005, 2006; Wu and Xia, 2006), and cara鄄
gana korshinskii (He et al., 2009a). In general,
AM plants show higher stomatal conductance, tran鄄
spiration rates, hydraulic conductivity and leaf water
potential under water stress conditions, indicating
that AM plants maintained more normal water rela鄄
tions ( Auge, 2004 ). However, the mechanisms
by which AM fungi have enhanced the water rela鄄
tions of host plants are still unclear. Potential mecha鄄
nisms mainly include enhanced absorption of water
by external hyphae (Ruiz鄄Lozano and Azcon, 1995),
stomatal regulation through hormonal signals (Goico鄄
echea et al., 1997 ), indirect effect of improved
phosphorus (Fitter, 1988), and a greater osmotic
adjustment in AM plants (Ruiz鄄Lozano, 2003).
Sod culture in orchard not only can effectively
avoid water and soil erosion, but also can enhance
organic matter, mineral content in soil ( Li et al.,
2007), hence, it is thought to be an important
planting technology in modern ecotype orchard.
Presently, bahia grass (Paspalum notatum Flugge)
is widly applied to sod culture in orchard of China
(Li et al., 2005, 2007), which can increase fruit
tree (such as citrus plant) root colonization by ar鄄
buscular mycorrhizae, and improve nutritional level
and fruit quality (Zeng et al., 2004; Li and Ying,
2005; Ren et al., 2009). Whereas, information a鄄
bout the effects of AM fungi on growth, osmotic ad鄄
justment and reactive oxygen metabolism of common
grass of sod culture in orchard like baiha grass is
scarce. The purpose of this study is to, evaluate the
effects of Glomus mosseae on growth, osmotic active
solutes and antioxidant enzymes of bahia grass under
water stress conditions, the result will play a theore鄄
tical and basic role taking practical steps for produc鄄
tion in the area.
Materials and methods
Experimental materials and growth conditions
Seeds of bahia grass were surface鄄sterilized with
70% alcohol for 5 min and germinated on wet filter
paper in Petri dishes in darkness at 25益 . Seven
days after germination, seedlings were transplanted
to a plastic pot containing 3. 0 kg of an autoclaved
growth substrate (0. 11 MPa, 121益, 2 h) of yellow
soil and quartz sand (9 颐 1, v / v), the characteris鄄
tic of which were: pH 6. 1, 1. 2% organic matter,
available P 11. 05 mg·g-1, available K 39. 28 mg·
g-1, available Ca 394. 07 mg·g-1, available Mg
11. 33 mg·g-1, available Zn 0. 47 mg·g-1, availa鄄
ble Cu 0. 09 mg·g-1 . The soil was collected from
the Experimental Sample Garden, Anhui Normal U鄄
niversity (Wuhu, China), The experimental pots
were placed in a greenhouse under natural light con鄄
ditions without any temperature control from April to
June. The inoculated and non鄄inoculated bahia grass
were irrigated once every 2 day. The average day /
night temperature was 30 / 25益 during water stress,
and average air relative humidity was 80% . AM and
non鄄AM plants were not fertilized during the entire
experiment.
Mycorrhizal fungus inocula
Mycorrhizal fungus inocula, consisting of spores,
soil, hyphae and infected root fragments from a stock
culture of Glomus mosseae (No. BGC HUN01A), were
provided by Bank of Glomales in China, Institute of
Plant Nutrition and Resources, Beijing Academy of
Agriculture and Forestry Sciences. The inoculated
dosage was 20 g of inocula per pot containing ap鄄
proximately 1488 spores. Mycorrhizal inocula were
placed 5 cm below baiha grass roots at the time of
transplanting. Non鄄AM treatments received the same
mass of autoclaved growth substrate.
Experimental design
The present experiment was a completely ran鄄
225摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
domized 2伊2 factorial design, with two mycorrhizal
treatments, AM fungus and non鄄AM fungus, and
four water treatments, cut off water supply for 0 day,
4 days, 8 days, and water supply restored for 4
days. Water stress treatment began after 75 days
(June 15, 2009) of acclimation in greenhouse con鄄
ditions. Each of the eight treatments was replicated
four times, leading to a total of 32 pots.
Plant growth and mycorrhizal colonization mea鄄
surement
At the time of water stress treatment began, A
fraction of the plants were harvested and plant
height, fresh weight of shoots and roots were recor鄄
ded. The shoots and roots were separated and dried at
75益 for 48 h. A fraction of fresh roots were carefully
washed and cut into 1 cm root pieces to fix in forma鄄
lin鄄acetic acid鄄alcohol solutions, these roots were
cleaned with 10% (w / v) KOH and stained with
0. 05% (w / v) trypan blue in lactophenol (Phillips
and Hayman, 1970), and microscopically observed
for root colonization. The AM infected percentage
was counted by the following formula: AM infected
percentage (% ) = 100 伊 root length infected / root
length observed.
Inorganic ions analyses
Hundred milligrams of dry shoots or roots were
used to extract inorganic ions with 20 mL 1 mol·L-1
hydrochloric acid at 40益 for 2 h, and then filtered
through a piece of filter paper. Phosphorus ( P),
potasium (K), calcium (Ca), magnesium (Mg),
iron (Fe), manganese (Mn), zinc (Zn), boron
(B) and copper (Cu) concentrations in the dried
shoots and roots were determined by inductively cou鄄
pled plasma optical emission spectrometry ( ICP鄄
OES, Iris鄄Advantage type, USA).
Relative water content, chlorophyll content and
relative conductivity determination
The equation relative water content ( RWC)
(% )= 100 伊 ( FW鄄DW) / ( SW鄄DW) was used for
RWC calculation, where FW stood for fresh weight,
DW for dry weight, and SW for saturated weight, SW
was determined after floating the shoot on distilled
water for 24 h at the room temperature. Shoot chloro鄄
phll content and relative conductivity were assayed
according to Li (2003). The extraction was made
from 50 mg鄄fresh sample in 25 mL acetone (80% )
in the dark at the room temperature and was mea鄄
sured at 646 and 663 nm with a UV / VIS spectropho鄄
tometer. Relative conductivity was determined by
conductivity meter (DDS鄄307).
Determination of SOD and POD acitivities, MDA
and proline contents
Another fraction of the plants were harvested at
the water treatment time (0 d, 4 d, 8 d and 12 d),
Samples were quick freezed by liquid nitrogen, and
stored in refrigerator for superoxide dismutase
(SOD), peroxidase (POD) activities, malondialde鄄
hyde (MDA) and proline contents analysis. Extracts
for determination of SOD, POD were prepared from
half a gram of fresh shoot tissue homogenized in 5
mL of 0. 1 mol·L-1 phosphate buffer (pH 7. 8) con鄄
taining 0. 1 mmol·L-1 EDTA, 1 mmol·L-1 ASC, 1
mmol·L-1 1,4鄄dithiothreitol and 2% (w / v) polyvi鄄
nylpyrrolidone and centrifuged at 4 200 g for 10 min,
with the resulting supernatant used for assays. All
steps of the extraction procedure were carried out at
4益 . SOD activity was measured using the method of
Giannopolitis and Ries (1977). One unit of SOD
was defined as the amount of enzyme that inhibited
50% nitro blue tetrazolium by light, and SOD activi鄄
ty was expressed as SOD units per mg of protein.
POD activity was determined using the method of
Chance and Maehly (1955). Five hundred mg of
fresh shoots were homogenized in 5 mL of 5% (w /
v) trichloroacetic acid and centrifuged at 3 000 g for
10 min. MDA of extracts was determined by the
thiobarbituric acid reaction as described by Sudhakar
et al. (2001).
For proline determination, fresh samples were
extracted with 3% sulfosalicyclic acid, placed in a
boiling water bath for 10min and filtered through fil鄄
ter paper. Two milliliter of extract was added to 6
mL assay media containing 2 mL ninhydrin solution
and 2 mL acetic acid and incubated for 30 min at
3255 期摇 摇 XIAO Jia鄄Xin et al. : Effects of Arbuscular Mycorrhizal Fungi on Physiological Character of Bahia Grass …摇 摇
100益 and then cooled. The colored product formed
was extracted with 4 mL toluene by shaking and the
absorbance of resultant organic layer was measured
at 520 nm (Troll and Lindsley, 1955).
Statistical analysis
The experimental data were subjected to analy鄄
sis of variance (ANOVA) using the Statistical Anal鄄
ysis System (SAS 8. 1) software. For comparison of
the means, the Duncan爷s multiples range test was
employed.
Results and analysis
No mycorrhizal colonization was observed in the
roots of non鄄AM plants. The roots of plants inocula鄄
ted with Glomus mosseae were infected. AM plants
had significant plant height, shoot and root fresh
weight than corresponding non鄄AM plants ( Table
1).
Phosphorus, K contents in shoots, P, Ca con鄄
tents in roots were higher in AM plants than those in
non鄄AM plants, and the P, K levels were notably
higher in shoots than in roots of both AM and non鄄
AM plants, whereas, AM colonization did not affect
the Mg contents in shoots and roots (Table 2).
The contents of Mn in shoots and roots of AM
plants were significantly higher than those in non鄄
AM plants, while shoot Zn content, root Fe, B, Cu
contents were substantially lower compared with cor鄄
responding non鄄AM plants, Fe contents were higher
in roots than in shoots of AM and non鄄AM plants
(Table 3).
As the water stress progresses, shoot relative
water content was decreasing degressively in non鄄AM
plants, whereas, changed slightly in AM plants.
During water stress, relative water contents were sig鄄
nificantly higher in AM than in non鄄AM plants (Fig.
1: A).
Chlorophyll contents of shoots were decreasing
degressively in non鄄AM and AM plants with the in鄄
crease of drought degree, and shoot chlorophyll con鄄
tents were higher in AM than in non鄄AM plants at
the stage of water stress (Fig. 1: B).
Table 1摇 Root and shoot characteristics of AM and non鄄AM Paspalum notatum plants
AM fungus status Root colonization (%) Plant height (cm) Leaf fresh weight (g·plant-1) Root fresh weight (g·plant-1)
non鄄AM fungus 0. 00a 23. 33依3. 06a 0. 25依0. 02a 0. 11依0. 02a
AM fungus 33. 87依5. 16b 41. 67依2. 89b 0. 49依0. 04b 0. 45依0. 04b
Data are Means依SD. the same letters within a column indicate no significant differences at P<0. 05. The same is in the following tables
Table 2摇 Phosphorus, K, Ca and Mg concentrations in shoots and roots of AM and non鄄AM Paspalum notatum plants
AM fungus
status
P (mg·g-1 DW)
Shoot Root
K (mg·g-1 DW)
Shoot Root
Ca (mg·g-1 DW)
Shoot Root
Mg (mg·g-1 DW)
Shoot Root
non鄄AM
fungus 1. 18依0. 19a 0. 52依0. 10a 23. 16依0. 11a 12. 57依0. 83a 8. 70依0. 15a 5. 58依1. 14a 2. 47依0. 38a 2. 44依0. 44a
AM
fungus 1. 95依0. 01b 0. 83依0. 02b 30. 99依4. 34b 13. 47依2. 13a 10. 16依1. 71a 9. 00依0. 66b 2. 65依0. 17a 3. 06依0. 25a
Table 3摇 Zinc, Fe, B, Cu and Mn concentrations in shoots and roots of AM and non鄄AM Paspalum notatum plants
AM fungus
status
Zn
(mg·kg-1 DW )
Shoot Root
Fe
(mg·kg-1 DW )
Shoot Root
B
(mg·kg-1 DW )
Shoot Root
Cu
(mg·kg-1 DW )
Shoot Root
Mn
(mg·kg-1 DW )
Shoot Root
non鄄AM
fungus
27. 47
依4. 18a
30. 35
依3. 34a
85. 55
依12. 82a
420. 89
依94. 63a
22. 8
依0. 58a
25. 4
依2. 42a
21. 68
依2. 03a
31. 38
依6. 28a
17. 53
依2. 50a
24. 68
依3. 81a
AM
fungus
16. 95
依2. 19b
27. 91
依6. 08a
76. 78
依6. 81a
304. 27
依32. 77b
15. 07
依3. 94a
12. 76
依7. 59b
19. 93
依2. 16a
20. 62
依2. 08b
46. 70
依3. 00b
58. 75
依6. 24b
425摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
Fig. 1摇 Relative water content (A), chlorophyll content (B), and
relative conductivity (C) of shoots in AM and non鄄AM Paspalum
notatum plants subjected to water stress Note: NM, non鄄AM plants;
AM, AM plants. The same is in the following figure
摇 Relative conductivity of shoots in non鄄AM
plants markedly increased at the time of water stress
for 4 days and 8 days, but no significant difference
was observed on shoot relative conductivity in AM
plants during water stress compared with correspond鄄
ing water stress before (Fig. 1: C).
Shoot SOD activities in non鄄AM and AM
plants, shoot POD activities in AM plants were in鄄
creasing progressively as the water stress progresses
( Fig. 2: A, B), but no significant change was
found on shoot POD activities in non鄄AM plants,
and during water stress, shoot POD activities were
conspicuously higher in AM than in non鄄AM plants
(Fig. 2: B). MDA contents of shoots were signifi鄄
cantly higher in non鄄AM than in AM plants during
water stress (Fig. 2: C). No significant change was
observed on proline in shoots from non鄄AM plants
during the time of water stress, whereas, the shoot
proline contents obviously increased in AM plants,
and were significantly higher than those in non鄄AM
plants (Fig. 2: D).
Discussion
We observed the positive effects of AM fungus,
Glomus mosseae, on the growth, the biomass and P,
K, Ca contents of bahia grass (Table 1, 2), which
indicated that AM colonization enhanced P uptake by
roots and its upward transportation, leading to the
plants爷 increasing growth and the other elements
such as K, Ca absorption by roots. Similar results
have been reported for other plant species (Wu and
Zou, 2007; Kapoor et al., 2008; Miransari et al.,
2009a; He et al., 2009b). Phophorus is generally
believed to be removing poorly in the soil, the grand
hyphal net formed in the soil by AM can travel
across P deficiency area around the root, which can
enhance the contact with the soil and turn unavaila鄄
ble P around root into available P to roots, the im鄄
provement of P, K, Ca absorption by AM plants,
and the direct uptake and transport of the extraradi鄄
cal hypha outside the roots are all likely to increase
the P, K, Ca levels in plant, which is consistent
with the findings that the field grass cultivation can
effectively increase the contents of P, K in citrus
leaves (Li and Yi, 2005; Ren et al., 2009), and
we suggested that the grass such as bahia grass in or鄄
chard have highly developed root system and be easi鄄
ly infected by AM fungi, which is beneficial to the
soil mineral elements, and thus promote P, K, Ca
uptake by roots. Additionally, AM fungi inoculation
also significantly increased Mn contents in shoots
and roots of bahia grass, but notably reduced Zn in
5255 期摇 摇 XIAO Jia鄄Xin et al. : Effects of Arbuscular Mycorrhizal Fungi on Physiological Character of Bahia Grass …摇 摇
Fig. 2摇 Superoxide dismutase (SOD) (A) and peroxidase (POD) (B) activities, and malondialdehyde (MDA) (C) and proline
(D) contents of shoots in AM and non鄄AM Paspalum notatum plants subjected to water stress
shoots and Fe, B, Cu in roots (Table 3), which is
not all in accordance with previous investigation on
trifoliate orange (Wu and Zou, 2007), corn (Mir鄄
ansari et al., 2009a) and wheat (Miransari et al.,
2009b). Generally, AM colonization has an influ鄄
ence on the number of reducing bacteria by the min鄄
eral elements such as Mn in the rhizosphere soil as
well as the reducing power of the rhizosphere. How鄄
ever, there existed differences for different hosts in鄄
fected by different AM fungi varieties, namely, AM
symbiosis and its host has different selectivity and a鄄
daptability, for example, AM fungi inoculation pro鄄
moted iron uptake by trifoliate orange roots, while
this whole process may be affected by the soil pH
value (Wang and Xia, 2009 ), additionally, soil
moisture conditions ( Song et al., 2008 ), P level
(He et al., 2009a), or nitrogen level (He et al.,
2009b) all affected soil nutrient uptake by the roots
infected by AM. Moreover, there is a mutual benefi鄄
cial and antagonistic relationship among mineral ele鄄
ments in the soil and plant. Thus, the differences of
research findings mentioned above should not only
be due to the species and characters of AM fungi and
their hosts, but also attribute to the interaction be鄄
tween the mineral elements in the soil and plants.
Water content in plant tissue can reflect cell
water status, and chlorophyll is the most important
and effective pigment in photosynthesis, which can
demonstrate the assimilation ability in plant. This
research showed that during water stress, AM plants
have a stable relative water and chlorophyll con鄄
tents, which were higher than those in non鄄AM
plants (Fig. 1), this result showed that AM fungi
inoculation can release the influence of water stress
on water metabolism and photosynthesis in shoots,
and improve the physiological status in order to
strengthen the anti鄄drought stress ability.
Water stress can induce ROS, as a result of lip鄄
id peroxidation and production of MDA (Lacan and
Baccou, 1998). Our study proved that there was
625摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
less MDA and relatively conductivity levels in AM
bahia grass shoots, enhancing drought tolerance of
AM plants. When plants are subjected to water
stress, activities of a number of antioxidant enzymes
are enhanced in order to elimilate ROS (Ruiz鄄Loza鄄
no, 2003). Wu et al. (2006) reported that activi鄄
ties of SOD, POD and CAT were usually higher in
AM than in non鄄AM roots under well鄄watered and
water stress conditions. In our results, under water
stress conditions, SOD activities increased in AM
and non鄄AM plants, while POD activity and proline
content increased significantly in shoots from AM
plants and were higher than those in non鄄AM plants
(Fig. 2). It is well known that the important func鄄
tion of SOD is to remove superoxide free radicals so
as not to generate H2O2, which is then elimilated by
POD and CAT. As a result, AM inoculation can re鄄
move the accumulation of reactive oxygen caused for
the water stress by improving SOD and POD activi鄄
ties, and adjust cell osmotic potential to protect the
activities of enzyme systems by promoting the accu鄄
mulation of proline as well as P, K, Ca, with the
aim of reducing the damage caused by membrane
lipid peroxidation, which in return enhance the
drought tolerance of bahia grass.
In short, G. mosseae鄄inoculation had a positve
effect on growth, osmotic adjustment, and reactive
oxygen metabolism of bahia grass, increasing the con鄄
centrations of proline, P, K, and POD in shoots,
and decreasing the shoot relative conductivity and
MDA content. The result suggested that the benefit
of AM colonization under water stress conditions was
due to the enhancement of osmotic adjustment and
higher antioxidant enzymes, and AM fungi inocula鄄
tion would be a new way relieasing the problems for
drought existed in sod culture in orchard.
References:
Auge RM, 2004. Arbuscular mycorrhizae and soil / plant water rela鄄
tions [J] . Canadian Journal of Soil Science, 84: 373—381
Chance B, Maehly AC, 1955. Assay of catalases and peroxidases
[J] . Methods Enzymology, 2: 764—775
Fitter AH, 1988. Water relations of red clover Trifolium pratense L.
as affected by VA mycorrhizal infection and phosphorus supply
before and during drought [J] . Journal of Experimental Botany,
39: 595—603
Giannopolitis CN, Ries SK, 1977. Superoxide dismutase. 玉. Occur鄄
rence in higher plants [J] . Plant Physiology, 59: 309—314
Goicoechea N, Antolin MC, Sanchez鄄Diaz M, 1997. Gas exchange is
related to the hormone balance in mycorrhizal or nitrogen鄄fixing
alfafa subjected to drought [ J] . Physiologia Plantarum, 100:
989—997
He XL (贺学礼), Liu T (刘媞), An XJ (安秀娟) et al., 2009a.
Effects of AM fungi on the growth and drought resistance of Cara鄄
gana korshinskii under water stress conditions [J] . Acta Ecolog鄄
ica Sinica (生态学报), 29 (1): 47—52
He XL (贺学礼), Liu T (刘媞), Zhao LL (赵丽莉), 2009b.
Effects of inoculating AM fungi on physiological characters and
nutritional components of Astragalus membranaceus under differ鄄
ent N application levels [J] . Chinese Journal of Applied Ecology
(应用生态学报), 20 (9): 2118—2122
Kapoor R, Sharma D, Bhatnagar AK, 2008. Arbuscular mycorrhizae
in micropropagation systems and their potential applications
[J] . Scientia Horticulturae, 116: 227—239
Kramer PJ, Boyer JS, 1997. Water Relations of Plants and Soils
[M]. San Diego: Academic Press, 1—20
Lacan D, Baccou JC, 1998. High levels of antioxidant enzymes corre鄄
late with delayed senescence in nonnetted muskmelon fruits [J] .
Planta, 204: 377—382
Li GH (李国怀), Yin HL (伊华林), Xia RX (夏仁学), 2005.
Application of bahia grass (Paspalum notatum) to eco鄄agricul鄄
ture construction in Southern China [J] . Chinese Journal of Eco鄄
Agriculture (中国生态农业学报), 13 (4): 197—199
Li GH (李国怀), Yin HL (伊华林), 2005. Influence of sod cul鄄
ture on the soil water content, effect of soil nutrients, fruit yield
and quality in citrus orchard [ J] . Chinese Journal of Eco鄄Agri鄄
culture (中国生态农业学报), 13 (2): 160—163
Li HK (李会科), Zhang GZ (张广军), Zhao ZY (赵政阳) et al.,
2007. Effects of interplanting of herbage on soil nutrient of non鄄
irrigated apple orchard in the loess plateau [J] . Acta Hortcultu鄄
rae Sinica (园艺学报), 34 (2): 477—480
Li HS (李合生), 2003. Experimental principle and technique for
plant physiology and biochemistry [ M ]. Beijing: Higher
Edueation Press, 167—169
Miransari M, Bahrami HA, Rejali F et al., 2009a. Effects of soil
compaction and arbuscular mycorrhiza on corn ( Zea mays L. )
nutrient uptake [J] . Soil & Tillage Research, 103: 282—290
Miransari M, Bahrami HA, Rejali F et al., 2009b. Effects of arbus鄄
cular mycorrhiza, soil sterilization, and soil compaction on wheat
(Triticum aestivum L. ) nutrients uptake [J] . Soil & Tillage Re鄄
search, 104: 48—55
Phillips JM, Hayman DS, 1970. Improve procedures for clearing roots
and staining parasitic and vesicular鄄arbuscular mycorrhizal fungi
7255 期摇 摇 XIAO Jia鄄Xin et al. : Effects of Arbuscular Mycorrhizal Fungi on Physiological Character of Bahia Grass …摇 摇
for rapid assessment of infection [ J] . Transactions of the British
Mycological Society, 55: 158—161
Ren Q (任群), Xiao JX (肖家欣), Chen SL (陈世林) et al.,
2009. Influences of sod culture on mineral nutrition in citrus
leaves and fruit quality [J] . Chinese Agricultural Science Bulle鄄
tin (中国农学通报), 25 (24): 407—409
Ruiz 鄄Lozano JM, 2003. Arbuscular mycorrhizal symbiosis and allevi鄄
ation of osmotic stress, new perspectives for molecular studies
[J] . Mycorrhiza, 13: 309—317
Ruiz 鄄Lozano JM, Azcon R, 1995. Hyphal contribution to water up鄄
take in mycorrhizal plants as affected by the fungal species and
water status [J] . Physiologia Plantarum, 95: 472—478
Song HX (宋会兴), Peng YY (彭远英), Zhong ZC (钟章成),
2008. Photosynthetic responses of AMF infected and AMF鄄free
Bidens pilosa L. to drought stress conditions [J] . Acta Ecologi鄄
ca Sinica (生态学报), 28 (8): 3745—3751
Sudhakar C, Lakshmi A, Giridarakumar S, 2001. Changes in the an鄄
tioxidant enzymes efficacy in two high yielding genotypes of mul鄄
berry (Morus alba L. ) under NaCl salinity [J] . Plant Science,
161: 613—619
Troll W, Lindsley JA, 1955. A photometric method for the determina鄄
tion of proline [ J] . Journal of Biological Chemistry, 215:
655—660
Wang MY (王明元), Xia RX (夏仁学), 2009. Effects of arbuscu鄄
lar mycorrhizal fungi on growth and iron uptake of Poncirus trifo鄄
liata under different pH [ J] . Acta Microbiologica Sinica (微
生物学报), 49 (10): 1374—1379
Wu QS (吴强盛), Xia RX (夏仁学), Hu ZJ (胡正嘉), 2005.
Effects of arbuscular mycorrhiza on drought tolerance of Poncirus
trifoliata [J] . Chinese Journal of Applied Ecology (应用生态
学报), 16 (3): 459—463
Wu QS, Xia RX, Zou YN, 2006. Reactive oxygen metabolism in my鄄
corrhizal and non鄄mycorrihizal citrus (Poncirus trifoliata) seed鄄
lings subjected to water stress [ J] . Journal of Plant Physiolo鄄
gy, 163: 1101—1110
Wu QS, Xia RX, 2006. Arbuscular mycorrhizal fungi influence
growth, osmotic adjustment and photosynthesis of citrus under
well鄄watered and water stress conditions [J] . Journal of Plant
Physiology, 163: 417—425
Wu QS (吴强盛), Zou YN (邹英宁), 2007. Influence of AM fungi
on nutrient uptake in leaves of trifoliate orange seedlings subjec鄄
ted to water stress [ J] . Acta Botany Boreal鄄Occident Sinica
(西北植物学报), 27 (1): 0074—0078
Zeng M (曾明), Li DG (李道高), Xiong BQ (熊丙全) et al.,
2004. Effects of sod culture management in citrus orchardson
AM infection in citrus roots and on fruit quality [J] . Journal of
Southwest Agricultural University (西南农业大学学报), 26
(2): 105—107
825摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷