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ABSTRACT
The influences of arbuscular mycorrhizal (AM) fungus on growth, gas exchange, chlorophyll concentration, chlo-
rophyll fluorescence and water status of maize (Zea mays L.) plants were studied in pot culture under well-watered
and drought stress conditions. The maize plants were grown in a sand and black soil mixture for 4 weeks, and then
exposed to drought stress for 4 weeks. Drought stress significantly decreased AM colonization and total dry weight.
AM symbioses notably enhanced net photosynthetic rate and transpiration rate, but decreased intercellular CO2
concentration of maize plants regardless of water treatments. Mycorrhizal plants had higher stomatal conductance
than non-mycorrhizal plants under drought stress. The concentrations of chlorophyll were higher in mycorrhizal
than non-mycorrhizal plants under drought stress. AM colonization significantly increased maximal fluorescence,
maximum quantum efficiency of PSII photochemistry and potential photochemical efficiency, but decreased prima-
ry fluorescence under well-watered and droughted conditions. Mycorrhizal maize plants had higher relative water
content and water use efficiency under drought stress compared with non-mycorrhizal plants. The results indicated
that AM symbiosis alleviates the toxic effect of drought stress via improving photosynthesis and water status of
maize plants.
Keywords: chlorophyll concentration; chlorophyll fluorescence; gas exchange; maize; water status
Plant is constantly faced with environmental trial plants and fungi of the monophyletic phylum
stress factors of both biotic and abiotic origin. Glomeromycota (Schüssler et al. 2001). AM fungi
Drought stress is one of the most important abiotic are known to improve the mineral nutrition of
stress factors that limits plant growth and devel- colonized plants, they also increase their resistance
opment in many arid and semiarid regions of the to drastic environmental conditions, especially
world, and seasonal drought often occurs in the drought stress (Smith and Read 2008).
non-arid regions. Drought caused a great reduc- It has been demonstrated that AM symbiosis
tion in productivity and yield of crop, although affects the water relations of many plants. The
crop characteristics were associated with improved AM symbiosis often results in the change of wa-
capability under drought stress which allows crops ter movement into, through, and out of the host
to absorb more water, to reduce water loss rates, plants, with consequent effects on tissue hydration
and to maintain higher physiological activity. In and plant physiology. The contribution of mycor-
addition, drought caused by climate change and rhizal symbiosis to drought tolerance is due to a
water shortages will become more serious, which combination of physical, nutritional, physiological,
will drastically affect growth and survival of plants and cellular effects (Aroca et al. 2008, Miransari
and crops in many parts of the world. 2010). AM and non-AM plants often show differ-
Arbuscular mycorrhizal (AM) symbiosis is ent photosynthetic characters. Wu and Xia (2006)
formed between the majority (70–90%) of terres- confirmed that AM citrus seedlings had higher
Supported by the National Nature Science Foundation of China, Project No. 31000679, and by the Key Laboratory
of Mollisols Agroecology, Chinese Academy of Sciences.
Means followed by the same letter are not significantly different (P < 0.05)
AM colonization did not result in higher plant Pn and E than non-mycorrhizal plants under well-
height and dry weight. This observation may be watered and drought stress conditions (Figure 1).
related to carbon drain effect or dysfunction of The g s in AM-inoculated plants was significantly
maize metabolism (Liu et al. 2004). higher than in non-inoculated plants under drought
The rate of gas exchange is an important factor stress. Compared with non-mycorrhizal plants,
that influences plant growth under drought stress. g s was 24.6% higher in mycorrhizal plants. The
Mycorrhizal plants often display higher P n than results implied that AM symbioses could enhance
non-mycorrhizal plants, which is consistent with the gas exchange capacity via decreased stomatal
AM symbioses effects on gs (cf. Augé 2001). In the resistances and increased transpiration fluxes. The
present study, mycorrhizal maize plants had higher lower values of stomatal resistance in mycorrhizal
14 M+ 180
a a a
12 M-– 160
b 140
gs (mmol H2O/m2/s)
10 b
Pn (µmol CO2/m2/s)
c 120
8 c
d 100
6 80
60
4
40
2 20
0 0
Well-watered Droughted Well-watered Droughted
3
140 a
a
2.5 b 120
E (mmol H2O/m2/s)
c
Ci (µmol CO2/mol)
100 b b
2 d
80 c
1.5
60
1
40
0.5
20
0 0
Well-watered Droughted Well-watered Droughted
Figure 1. Net photosynthetic rate (P n), transpiration rate (E), stomatal conductance (g s) and intercellular CO 2
concentration (Ci) of maize plants inoculated (M+) or not (M–) with Glomus etunicatum under well-watered
and drought stressed conditions. Mean pairs followed by different letters are significantly different (P < 0.05)
Means followed by the same letter are not significantly different (P < 0.05)
plants indicate that these plants were able to keep Chlorophyll fluorescence was proven to be a
the stomata open longer than non-mycorrhizal very useful noninvasive tool for the evaluation of
plants (Subramanian et al. 1995). Some studied environmental stresses effect on photosynthetic
suggested that AM colonization may increase the properties. In the present study, the primary fluo-
number of photosynthetic units, and the rates of rescence (Fo) of mycorrhizal plants was lower than
photosynthetic storage and export (cf. Augé 2001). that non-mycorrhizal plants under drought stress
Chlorophyll status is a key index for evaluating (Table 2). Mycorrhizal plants had markedly higher
plant photosynthetic efficiency and environmen- maximal fluorescence (F m) than non-mycorrhizal
tal stress. The concentrations of chlorophyll a, plants regardless of water treatments. Drought
chlorophyll b and chlorophyll a + b were similar stress increased Fo, but decreased Fm of both AM-
in mycorrhizal and non-mycorrhizal maize plants inoculated and non-inoculated plants. The results
under well-watered conditions (Table 2). However, indicated that drought stress destructs the structure
the differences of chlorophyll a, chlorophyll b and and function of PSII reaction center and disrupts
chlorophyll a + b concentrations were significant electron transport in photosynthetic apparatus,
between AM and non-AM maize plants under whereas AM symbiosis mitigates the adverse in-
drought stress conditions, which is in agreement fluence of drought stress on PSII reaction center
with the results of other studies (cf. Augé 2001, and photosynthetic efficiency (Baker 2008).
Gemma et al. 1997). Compared with non-mycor- The ratio of Fv/Fm is a useful relative measure of
rhizal plants, the concentrations of chlorophyll a, the maximum quantum efficiency of PSII photo-
chlorophyll b and chlorophyll a + b of mycorrhizal chemistry, which provides a simple and rapid way
plants were 18.6%, 27.5% and 20.5% higher under of monitoring environmental stress (Krause and
drought stress, respectively. This suggests that Weis 1991, Baker 2008). In this study, the F v/Fm in
drought stress interferes less with chlorophyll mycorrhizal maize plants was higher than in non-
synthesis and/or more with chlorophyll breakdown mycorrhizal plants regardless of water treatments,
in mycorrhizal than in non-mycorrhizal plants and drought stress drastically decreased the ratio
(Evelin et al. 2009). of Fv/Fm (Figure 2), which is in agreement with the
0.81 M+ 4.5
a
0.8 M-– a
0.79 b 4
b
0.78
c
Fv/Fm
c
v 0
3.5
Fv/Fm
F /F
o
0.77
Fv/F
d
0.76 d 3
0.75
0.74 2.5
0.73
0.72 2
Well-watered Droughted Well-watered Droughted
Figure 2. Maximum quantum efficiency of PSII (F v/F m) and potential photochemical efficiency (F v/F o) of maize
plants inoculated (M+) or not (M–) with Glomus etunicatum under well-watered and drought stressed condi-
tions. Mean pairs followed by different letters are significantly different (P < 0.05)
75 3
b
70 2
65 1
60 0
Well-watered Droughted Well-watered Droughted
Figure 3. Relative water content (RWC) and water use efficiency (WUE) of maize plants inoculated (M+) or not
(M–) with Glomus etunicatum under well-watered and drought stressed conditions. Mean pairs followed by
different letters are significantly different (P <0.05)
results by Borkowska (2002). At the same time, 2004) than non-mycorrhizal plants. However, the
the parameter F v/F o, an alternative expression of RWC in mycorrhizal and non-mycorrhizal plants
F v/F m, is a more sensitive and better value than was similar under well-watered conditions, which
F v/F m (Maxwell and Johnson 2000), also higher in indicated that leaf RWC of non-stressed plants
mycorrhizal plants. These results indicated that were not affected by AM symbiosis.
AM symbioses might improve photochemistry ef- Water status of mycorrhizal plants was often asso-
ficiency of PSII under well-watered and droughted ciatied with gas exchange, such as stomatal behavior
conditions. and transpiration fluxes. Augé (2001) pointed out
It is well documented that AM symbiosis can that gs and leaf water potential are linked function-
improve the water status of host plant. Porcel and ally: changes in one usually drive changes in the
Ruiz-Lozano (2004) reported that mycorrhizal other. Higher RWC and WUE in mycorrhizal than
plants had higher leaf water potential compared in non-mycorrhizal plants may be beneficial for
with non-mycorrhizal plants under drought stress. moving water through the plants to the evaporat-
Our study showed that AM symbiosis enhanced ing surfaces and maintaining opened stomata in
WUE (by 7.0% and 11.8%) in maize plant leaves leaves (Nelsen and Safir 1982). Similarly, higher g s
under well-watered and droughted conditions and E in mycorrhizal compared to non-mycorrhzial
(Figure 3). Drought stress significantly decreased plants may indicate lower resistance to water vapor
WUE in both mycorrhizal and non-mycorrhizal transfer from inside the leaves to the atmosphere,
plants. AM colonization also affected RWC of which allows leaves to maintain more normal water
maize plants. Mycorrhizal plants had higher RWC balance (Augé 2004, Wu and Xia 2006). Thus, the
than non-mycorrhizal plants under drought stress, results suggested that AM symbiosis protects maize
but there was no difference between mycorrhizal plants against drought stress through improving
and non-mycorrhizal plants under well-watered gas exchange capacity, chlorophyll concentration,
conditions. Drought stress markedly decreased chlorophyll fluorescence and water status.
RWC of non-mycorrhizal plants, but caused no
significant difference between well-watered and
droughted conditions for mycorrhizal plants. REFERENCES
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through hormonal signals (Aroca et al. 2008), application are modulated differently by mycorrhization in
indirect effect of improved phosphate and other tomato and an ABA-deficient mutant (Sitiens). Microbial
nutrient uptake (Subramanian and Charest 1997), Ecology, 56: 704–719.
greater osmotic adjustment (Wu and Xia 2006) Augé R.M. (2001): Water relations, drought and vesicular-arbus-
and higher root hydraulic conductivity (Augé cular mycorrhizal symbiosis. Mycorrhiza, 11: 3–42.
Corresponding author:
Dr. Xiancan Zhu, Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences,
Changchun, P.R. China
e-mail: zhuxiancan@neigae.ac.cn