Screening of Faba Bean (Vicia faba L.) Genotypes for
Mycorrhizal Association Parameters
Gemechu Abu
1*, Christian A. Fatokun
2, Gemechu Keneni
3, Fassil Assefa
4and Zerihun Belay
51Madda Walabu University, P.O. Box 247 Bale-Robe, Ethiopia
2International Institute of Tropical Agriculture, P.O. Box 200001 Ibadan, Nigeria 3Holetta Agricultural Research Center, P.O. Box 31 Holetta, Ethiopia
4Addis Ababa University, P.O. Box 1176 Addis Ababa, Ethiopia
5Adama University, P.O. Box 1888 Adama, Ethiopia
Faba bean (Vicia faba L.) is one of the most important legumes used for food, feed and maintaining ecological balance. However, its productivity has been declining due to various biotic and abiotic factors. Arbuscular Mycorrhizal symbioses have been proved to enhance growth and yield responses of faba bean and other agricultural crops by counteracting these extreme factors. The study was conducted at field and greenhouse to evaluate the performance of faba bean genotypes for mycorrhizal colonization on two Phosphorus fertilizer levels. Randomized Complete Block Design with three replications was used. Analysis of variance indicated that all parameters, except number of mycorrhizal spores and relative mycorrhizal dependency, were highly significantly (p<0.01) varied for genotypes. Genotypes Wayu, Dosha, Didea, Moti and Tumsa for phosphorus fertilized; and Dagim, Gebelcho, Dosha, Tumsa and Wayu for unfertilized trial had performed better. Mycorrhizal colonization performance was higher on unfertilized than on P fertilized trial, at both field and greenhouse. Furthermore, for the genotypes tested at both conditions, the performance of the genotypes was higher at field than at greenhouse. The study also revealed that most of the mycorrhizal association parameters were highly correlated with biomass phosphorus uptake and grain yield of the faba bean genotypes.
Key words: arbuscular mycorrhiza, faba bean, screening, correlation
INTRODUCTION
Faba bean is one of the most important food legumes ranking fourth in the world, after field pea, chickpeas and lentil. The world area of faba bean production is 2.5 million ha (FAOSTAT, 2016). It is cultivated in the temperate and subtropical regions of the world (Maxted and Bennett, 2001; Torres et al., 2006). Ethiopia is the leading producer of faba bean in Africa with a share of 1.5 million metric ton in 2013 (FAOSTAT, 2016). The productivity of improved varieties is very high (3.5 t/ha) compared to the country average yield (1.8 t/ha) (Maalouf et al., 2017).
Despite the potential, there has been a steady reduction in the cultivated area of faba bean in many countries. Many reasons have been suggested for this general decline but the major constraint to the crop is lack of adapted cultivars resistant to the major biotic and abiotic factors (van Emden
et al., 1988). For low input agricultural systems of Ethiopia
where chemical fertilizers, particularly phosphorus fertilizers, are rarely used in the production of faba bean and other pulse crops cereal crops (Mulissa and Fassil, 2012), the magnitude of the problem is exacerbating. Fertilizer applied area under faba bean ranges between 21% in 2000/01 to 28% in 2010/11 cropping season CSA (2011). Hence, an in-depth look at the alternatives or means to overcome such problems needs to be examined in order to ensure sustainable production of the crop.
*Corresponding Author: Gemechu Abu, Madda Walabu University, P.O. Box 247 Bale-Robe, Ethiopia. E-mail:
gemechuabu2002@gmail.com; Tel: +251-912057788
Co-Author 2Email: C.FATOKUN@cgiar.org; 3Email: gemechukeneni@yahoo.com; 4Email: asefafasil2013@gmail.com; 5E-mail: zebelay2009@gmail.com
Research Article
It has been proved that growth and yield responses of agricultural crops were improved and reduction of inputs such as P fertilizer was achieved via manipulation of Arbuscular Mycorrhizae (AM) symbioses (Sawers et al., 2010, Saia et al., 2014). AM is the most widespread terrestrial symbiosis and about 90% of land plants are able to form AM symbiosis (Smith and Read, 2008). The plant provides the fungi with carbohydrates and, in exchange, the fungi increase the plant’s ability to take up phosphorus and micronutrient from the soil (Tawaraya, 2003), provide protection from diseases (Borowicz, 2001; Tanwar et al., 2013), and improve the plant’s ability to access water from the soil (Zhu et al., 2012). Huiying et al., 2006 indicated that over 50% of P uptake by wheat plants was absorbed via AM fungi, even when P was added. Under low N and P fertilizer inputs or lower amount of nutrients in the soil, AMF has contributed for increased rate of N2-fixation in legume crops (Andrade et al., 1998).
The symbiotic associations between faba bean and AM fungi have resulted in the crop’s increased biomass production and photosynthetic rates by increasing the rate of phosphorus and nitrogen accumulation (Jia et. al., 2004). For the experiment on FB with suboptimal soil nutrient contents, higher nutrient uptakes of faba bean were found after application of AMF in comparison to the control (Almethyeb et al., 2013).
AM colonization differed among plant genotypes (Tawaraya, 2003, Leiser et al., 2015). Zhu et al. (2001) reported smaller % AM colonization in modern varieties than landraces of wheat varieties; while Koide et al.,1988; Bryla and Koide,1990 reported the opposite for oat (Avena sativa) and tomato (Solanum esculentum). Toth et al. (1990) further reported that colonization decreased for diseases-resistant lines, compared to susceptible lines of maize (Zea mays) inbred lines. Even though variation in AM colonization among genotypes of faba bean has been reported in other countries, there is very limited information on performance evaluation of Ethiopian faba bean genotypes for mycorrhizal association. Hence, it’s hypothesized that the genotypes will be significantly different from one another for mycorrhizal association parameters. Thus, the study was conducted with the objective of evaluating the potential and level of the genetic variation of AM colonization among genotypes of Ethiopian faba bean genotypes for AM colonization on two phosphorus fertilizer levels.
MATERIALS AND METHODS
Description of the Study areas
The study was carried out in both field and greenhouse conditions from 2015 to 2017. The field experiments were conducted under rain-fed condition in two faba bean growing areas of Ethiopia namely Adadi and Holetta. A greenhouse experiment was conducted at Addis Ababa
University, Ethiopia. A detailed description of the study areas and their soil physicochemical properties are indicated in Table 1.
Table 1. General description of the study areas and their soil physico-chemical properties
Parameters Adadi Ambo (G) Holetta
Altitude (masl) 2520 ---- 2390
Latitude (N) 8.21 ---- 9.04
Longitude (E) 38.29 ---- 38.03
Temperature (OC) 8.5-23.5 ---- 6.4 -24.4
Rainfall (mm) 930.8 ---- 760.8
Soil type Vertisol Vertisol Nitisol
Soil textural class Clay Clay Clay
% Clay 61.18 66.58 46.42
% Silt 25.34 15.25 32.48
% Sand 12.54 15.45 20.17
pH (H20) 6.4 6.79 7.3
Available P (ppm) 15.94 19.92 23.67
Total N (%) 0.15 0.17 0.18
K (ppm) 37.35 31.56 25.79
Organic C (%) 1.16 1.17 0.738
CEC (Meq/100g) 25.13 18.17 23.05
EC (μS) 405.63 -- 697.67
Note: G=greenhouse
Experimental Design, Materials and Setup
Randomized Complete Block Design (RCBD) with three replications was applied to establish the field and greenhouse experiments. Twenty faba bean (Vicia faba) genotypes for field and twelve genotypes for greenhouse trials were obtained from Holetta Agricultural Research Center. The number of genotypes for greenhouse trial were reduced because of lack of sufficient greenhouse space. The number was set to twelve based on the preliminary information of the genotypes for their resistance or susceptibility to major disease and pest of the crop. The genotypes were highly commercialized high yielding varieties and most promising lines.
Mycorrhizal Association
To establish mycorrhizal association trial on the field, the occurrence of naturally available mycorrhizal fungi in the rhizosphere of the plant was considered; and thus, there was no artificial inoculation of the mycorrhizal fungi or spores to the experimental plots. Leiser et al., 2015 also conducted vast mycorrhizal research on the un-inoculated field.
For greenhouse mycorrhizal trial, the mycorrhizal inoculum was obtained from Ziway area, Southeast Ethiopia, from the rhizosphere of ten acacia trees growing at an average distance of 30m from one another. The rhizosphere soil of the trees in the specified site was confirmed to have high and sufficient amount of AMF inoculum density to establish a mycorrhizal association. Three replicates of 500 g of the rhizosphere soil were taken into a depth of 30 cm under each tree (Belay et al., 2013). The soil was composited and mixed well to have uniform distribution or number of mycorrhizal spore per unit weight of the soil inoculum. 200g of the soil was added to each pot as a source of mycorrhizal spore inoculum and mixed well with the upper 15 cm soil depth of each pot. Non-mycorrhizal pots received 200g of oven dry soil inoculum containing no mycorrhizal propagules (Gazey et al. 1992). Acaulospora spp and Funneliformis spp were the major species of the mycorrhizae recovered from Ziway. The fungi were chosen due to well-characterized capabilities in spreading external hyphae and P uptake (Belay et al., 2013). Schweiger et al., 2007 had reported that both fungi increased plant growth and phosphorus uptake.
The spore density of mycorrhizal fungi was determined twice: before sowing and after crop harvesting. AMF fungal spores were extracted from soil by Wet-Sieving and Decanting, as described by (Gerdemann and Nicolson, 1963), and by Sucrose Centrifugation, as described by Smith and Skipper (1967). 40 fields of observations were counted and the average was recorded as the spore density of mycorrhizal fungi per 100g of soil.
In order to assess or quantify the extent of mycorrhizal association or colonization in a root sample, the roots were first stained by the procedure developed by Phillips and Hayman, (1970) and the roots were made ready for observation. The stained root fragments were spread in petri plates with grid lines on the bottom so that the level of infection was determined using the grid line intersect method (Giovannetti and Mosse, 1980) under dissecting microscope at 40x magnification. Each intersection of root and gridline was checked for the presence or absence of AMF structure(s) and scored as colonized or not colonized by AMF. Using these values, the percentage of AMF colonization was calculated. 100 root-gridline intersects were considered for best representation (Brundrett et al., 1996).
Soil and Plant Analysis
Soil physico-chemical analysis was conducted for soil
samples (0-30cm), at Addis Ababa University
Ecophysiology laboratory and Hawassa University Plant and Soil Laboratory, based on the procedure described by Sahelemedihin and Taye (2000).
DATA COLLECTION
Mycorrhizal Association Parameters
Mycorrhizal Spore Number or Density: spore number or density per 100g of soil was determined by Wet-sieving and Decanting method (Gerdemann and Nicolson, 1963) followed by Sucrose Centrifugation, as described by Smith and Skipper (1967).
Percentage of mycorrhizal colonization (PMC): was determined using the grid-line intersection method (Giovannetti and Mosse, 1980) and calculated by the formula:
PMC = 𝐍𝐨.𝐨𝐟 𝐫𝐨𝐨𝐭 𝐬𝐞𝐠𝐦𝐞𝐧𝐭𝐬 𝐢𝐧𝐟𝐞𝐜𝐭𝐞𝐝
𝐓𝐨𝐭𝐚𝐥 𝐧𝐨.𝐨𝐟 𝐫𝐨𝐨𝐭 𝐬𝐞𝐠𝐦𝐞𝐧𝐭𝐬
𝒙 𝟏𝟎𝟎
Two more mycorrhizal parameters, namely, mycorrhizal Phosphorus uptake response and relative mycorrhizal dependency were measured for greenhouse trial. The parameters were not measured for field trial because it’s difficult to maintain the control status of mycorrhizal treatment at field condition.
Mycorrhizal P uptake response (MPR): was calculated according to Cavagnaro et al. (2003), using P uptake values of mycorrhiza inoculated plants and P uptake value of mycorrhiza un-inoculated plant as;
MPR = [(Mp - NMp)/NMp] *100,
where, MP is a plant inoculated with mycorrhiza and NMP is a plant which is not inoculated with mycorrhiza
Relative mycorrhizal dependency (RMD): was
calculated by expressing the difference between shoot dry weights of mycorrhizal (SDWM+) and non-mycorrhizal
plant (SDWM-) as a percentage of the shoot dry weight of
the mycorrhizal plant (Waceke et al., 2001). RMD = [(SDWM+ - SDWM-)/ SDWM+] *100
Biomass P uptake, BPU was calculated as a product of above ground dry biomass weight and plant phosphorus concentration.
Grain yield, GY was collected from 5 and 3 plants for field and greenhouse experiments, respectively.
Relative reduction of the performance of phosphorus untreated to the respective phosphorus treated plants was calculated to evaluate the sensitivity of the characters to phosphorus unavailability as
Statistical Data Analysis
Data were checked for homogeneity of variance and transformed, where applicable, to normalize the data before statistical analysis. An individual site and combined analysis of variance were performed using SAS 9.3 (SAS Institute, 2012). Multiple mean comparisons were performed using Tukey’s HSD test at 0.05 level of probability. Pearson’s phenotypic correlation coefficients were estimated using the PROC CANCORR subprogram of SAS.
RESULTS AND DISCUSSIONS
Mycorrhizal Association performance
Number of mycorrhizal spore (NMS): Analysis of variance indicated that the number of mycorrhizal spore was not significantly varied for different genotypes at all variants of the experiment; except for phosphorus fertilized greenhouse trial (Table 2).
Generally, genotypes performance for NMS per 100g of soil ranged from 509.34 to 575.43 on fertilized field trial; while the range was from 557.71 to 605.55 on unfertilized trial. At the greenhouse, it ranged from 330.91 to 455.66 on fertilized; while from 414.24 to 516.59 on unfertilized plots (Figure 1). Performance of the genotypes for NMS was -11.06% lower at the field; and -14.70 lower at the greenhouse. Kucey and Paul, 1983 had reported lower values of NMS for faba bean, which is certainly due to environmental variation. Furthermore, Kucey and Paul, 1983 showed that the number of spores extracted from soils fertilized with 45 kg P ha-1 was less than half the number of spores extracted from nonfertilized soils, indicating the suppressive effect of Phosphorus on number of mycorrhizal spores.
Mycorrhizal Colonization Percentage (MCP): Highly significant (p<0.001) variation was observed among genotypes for mycorrhizal colonization percentage at both field and greenhouse; and for both phosphorus levels (Table 2). Reports by various authors on different crops have indicated that there was large genotypic variation for mycorrhizal colonization percentage (Galvan et al. 2007; Tawaraya 2003; An et al.2010; Hildermann et al., 2010; Singh et al., 2012). Hajiboland et al., 2009 reported that genotypic difference in responsiveness to inoculation with AMF is attribuTable to the different contribution of mechanisms for increased nutrient uptake in mycorrhizal plants depending on nutrient, nutritional status and nutrient use efficiency of genotypes.
At field condition, Hachalu, followed by Tumsa, Moti, and Gebelcho on phosphorus fertilized plots; Dosha, followed by Wayu, Moti and Tumsa on unfertilized trial had the highest mycorrhizal colonization percentage (Figure 2). Under greenhouse condition, Wayu, followed by
Gora, and Dosha for fertilized trial; Wayu, followed by ILB4358, and Selale for unfertilized trial were the top performing genotypes for mycorrhizal colonization percentage (Figure 2). Jia et al., 2004 and Mohamed et al., 2013 reported similar values of MCP for faba bean. Performance of the genotypes for MCP on P fertilized was -17.50% lower at the field; and -27.66 lower at the greenhouse than on unfertilized trial. Our result is supported by the report of Kucey and Paul, 1983 and Leiser et al., 2015 who indicated that AM colonization was significantly higher in low-P than high-P conditions.
Mycorrhizal Phosphorus uptake response (MPR): Faba bean genotypes were highly significantly (p<0.001) different for MPR, for both phosphorus regimes (Table 4). Moti (70.35%), Hachalu (69.14%) and Didea (65.23%); and Hachalu (90.93%), Dosha (84.61%) and Moti (84.23%) had the highest MPR performance on fertilized and unfertilized trial respectively.as compared to other genotypes (Table 4).
Relative mycorrhizal dependency (RMD): Analysis of variance indicated that RMD performance of the genotypes was not significantly varied for phosphorus fertilized; while highly significant (p<0.001) variation among genotypes were observed for unfertilized trial. (Table 2). Wayu (71.78%), Didea (69.29%) and ILB4358 (68.49%) had the top three relative mycorrhizal dependency performance. (Table 4). The higher values of RMD are supported by the work reported by Plenchette et al. (1983) which indicated that legumes including faba bean have a high mycorrhizal dependency.
Biomass phosphorus uptake (BPU): Highly significant (p<0.001) variation was observed among genotypes for BPU performance. (Table 2). Nebiyu et al., 2016 had also reported that faba bean genotypes vary for their phosphorus uptake response. As shown on Table 3, genotypes Hachalu, Didea, Dosha, Tumsa, and Moti on phosphorus fertilized field trial; Moti, Dosha, Walki, Tumsa and Hachalu on unfertilized field trial had the highest amount of BPU. For greenhouse trial, Hachalu, Dosha and Moti were the best performing genotypes; on both fertilized and unfertilized trial (Table 4). Genotypes with higher mycorrhizal colonization had higher BPU on fertilized trial as opposed to the ones on unfertilized trial. This could be explained by the fact that in low p soil, primary root-induced P-uptake was shown to be down-regulated by the mycorrhizal uptake which can account for more than 80 % of the total P acquisition by the host plant (Smith et al., 2003 and Smith and Smith 2012).
Table 2. Descriptive statistics and relative reductions of mycorrhizal association parameters
Char. With Phosphorus Without Phosphorus RR
Mean SD MSG CV Mean SD MSG CV
Field
NMS 527.64 67.13 0.00ns 2.05 586 66.78 0.00ns 1.82 -11.06
MCP 41.88 5.13 102.48** 12.73 49.21 7.06 145.80** 14.87 -17.5
BPU 0.52 0.09 0.00** 17.06 0.33 0.05 0.00** 14.84 36.59
GY 73.01 4.17 33.78*** 5.89 62.96 3.71 43.80*** 5.71 13.76
Greenhouse
NMS 408.39 54.96 0.01** 2.31 468.44 55.82 0.00ns 1.95 -14.7
MCP 35.43 3.63 28.05** 10.64 45.23 2.98 15.03* 8.86 -27.66
MPR 58.64 8.48 131.97** 14.45 76.2 10.15 222.54** 13.32 -29.95
RMD 46.59 8.95 107.49ns 15.21 61.46 10.62 237.64** 17.29 -31.92
BPU 0.26 0.01 0.00** 5.31 0.16 0.01 0.00** 5.82 37.27
GY 37.75 2.52 30.57*** 6.67 30.41 2.12 25.49*** 6.97 18.88
Note: NMS (per 100g soil), Number of mycorrhizal spore; MCP (%), Mycorrhizal colonization percentage, MPR, Mycorrhizal Phosphorus uptake response; RMD, Relative mycorrhizal dependency; BPU, Biomass phosphorus uptake; GY, Grain Yield
Table 3. Effect of mycorrhizal colonization on phosphorus uptake and grain yield of faba bean genotypes
Genotypes With Phosphorus Without Phosphorus
BPU (g/kg) GY (g/5p) BPU (g/kg) GY (g/5p)
Lalo 0.481fg 71.31b-e 0.298ef 61.23b-d
Dagim 0.527b-e 72.42a-e 0.333b-d 62.67b-d
EH06088-1 0.464g 71.03c-e 0.293f 59.01d
CS20DK 0.515c-f 74.24a-e 0.325c-e 63.21b-d
Obse 0.525b-f 73.56a-e 0.334b-d 61.85b-d
Gebelcho 0.533b-d 78.31a 0.333cd 64.80a-d
Holetta-2 0.506c-g 73.46a-e 0.318c-f 63.20b-d
Hachalu 0.580a 78.21ab 0.346a-d 62.65b-d
Wayu 0.528b-e 71.72a-e 0.308ef 59.94cd
Selale 0.484e-g 68.55e 0.322c-e 62.45b-d
Didea 0.564ab 72.93a-e 0.339b-d 63.83a-d
Gora 0.511c-f 72.22a-e 0.341a-d 64.32a-d
Dosha 0.551a-c 75.95a-d 0.353ab 66.34ab
EH07015-7 0.526b-f 71.62a-e 0.333cd 62.47b-d
EH06022-4 0.504d-g 69.19de 0.344a-d 60.06cd
Walki 0.530b-d 73.80a-e 0.350ab 65.14a-c
NC58 0.507c-g 70.73c-e 0.313c-f 61.35b-d
Moti 0.543a-d 76.83a-c 0.364a 69.19a
Tumsa 0.546a-d 73.63a-e 0.353ab 64.34a-d
EH06006-6 0.507c-g 70.51c-e 0.321c-e 61.16b-d
Mean 0.522 73.01 0.331 62.96
LSD 0.046 6.85 0.032 5.88
Mean Sq. 0.00** 33.78*** 0.00** 43.80***
CV 17.06 5.89 14.84 5.71
phosphorus levels (Table 3). For greenhouse trial, at both phosphorus fertilized and unfertilized trial, Moti and Dosha had the highest grain yield; while Tumsa and Walki had the lowest grain yield performance (Table 4).
As observed in Table 3 and 4, performances of the genotypes for grain yield were markedly varied at two
Screening of Faba Bean (Vicia faba L.) Genotypes for Mycorrhizal Association Parameters
Table 4. Effect of mycorrhizal colonization on mycorrhizal phosphorus uptake response, relative mycorrhizal dependence, phosphorus uptake and grain yield of faba bean at greenhouse
Genotypes With Phosphorus Without Phosphorus
MPR RMD BPU GY MPR RMD BPU GY
Obse 54.22ab 39.90a 0.268a 37.99c 65.22bc 53.08ab 0.164a 29.46cd
Hachalu 69.14a 55.50a 0.264ab 41.73ab 90.93a 66.96ab 0.162ab 34.38ab
ILB4358 60.25ab 50.06a 0.256ab 37.29cd 76.03a-c 68.49a 0.159ab 31.36bc
Selale 56.35ab 53.96a 0.255ab 34.30de 71.53a-c 65.09ab 0.159ab 28.71de
Didea 65.23ab 45.22a 0.259ab 39.75bc 84.10ab 69.29a 0.160ab 32.37bc
Gora 61.75ab 52.28a 0.260ab 40.19b 75.36a-c 65.72ab 0.162ab 31.84bc
Dosha 64.25ab 48.09a 0.260ab 43.28a 84.61ab 65.50ab 0.161ab 35.37a
Walki 50.65b 35.02a 0.256ab 36.84c-e 66.93bc 57.31b 0.163ab 28.05de
Moti 70.35a 40.90a 0.251bc 43.78a 84.23ab 45.19b 0.162ab 35.86a
Tumsa 51.85b 45.86a 0.245bc 34.06e 63.18c 46.53b 0.155b 27.49e
Gebelcho 52.15b 46.11a 0.242c 41.32ab 75.00a-c 62.60ab 0.153b 34.65ab
Wayu 51.45b 46.24a 0.250bc 37.93c 77.28a-c 71.78a 0.161ab 30.48cd
Mean 58.75 46.59 0.256 37.75 76.2 61.46 0.160 30.41
LSD 17.12 21.21 0.012 3.21 20.42 25.13 0.007 3.01
MSG 131.97** 107.49ns 0.00** 30.57*** 222.54** 237.64** 0.00** 25.49***
CV 14.45 19.21 5.31 6.67 13.32 17.29 5.82 6.97
Note: LSD, least significant difference; MSG, mean square of genotype; CV, coefficient of variation
a)
b)
c)
d)
Screening of Faba Bean (Vicia faba L.) Genotypes for Mycorrhizal Association Parameters
a) b)
c) d)
Figure 2. Mycorrhizal colonization percentage of genotypes a) at P fertilized Holetta and Adadi fields, b) for P fertilized and unfertilized field trial, c) at P unfertilized Holetta and Adadi fields, d) for P fertilized and unfertilized greenhouse trial
Correlation among mycorrhizal colonization
parameters
As shown in Tables 5 and 6, mycorrhizal colonization percentage was highly correlated with grain yield at all test conditions and with biomass phosphorus uptake for fertilized field trial (r = 0.24) and fertilized greenhouse trial (r = 0.46). Schweiger et al., 2007 also reported that mycorrhizal root colonization increased shoot P contents except at the native P status of the soil and when the highest P rate was applied. At both field and greenhouse of unfertilized trial, biomass phosphorus uptake was not correlated with mycorrhizal colonization percentage. Leiser et al., 2015 also reported a lack of correlation among the parameters at low phosphorus soil.
Mycorrhizal spore number had no correlation with mycorrhizal colonization percentage. It was highly correlated with grain yield at field trial as opposed to lack of correlation with the parameter at the greenhouse. At the greenhouse, relative mycorrhizal dependence and mycorrhizal phosphorus uptake responsiveness were strongly correlated to each other; suggesting an important contribution of shoot growth response to P uptake of mycorrhizal plants. This finding was also supported by the report of Hajiboland et al., (2009). However, the two characters did not correlate with mycorrhizal colonization percentage, except with relative mycorrhizal dependence on unfertilized trial.
Table 4. Correlation among mycorrhizal parameters at field
Char. P BPU NMS MCP GY
BPU P+ 1
BPU P- 1
NMS P+ .52** 1
NMS P- .37** 1
MCP P+ .24** 0.01 1
MCP P- 0.13 0.1 1
GY P+ .35** .20* .48** 1
GY P- .49** .24** .35** 1
Table 5. Correlation among mycorrhizal parameters at greenhouse
Char. P BPU NMS MCP MPR RMD GY
BPU P+ 1
BPU P- 1
NMS P+ 0.2 1
NMS P- 0.09 1
MCP P+ .46** 0.02 1
MCP P- 0.27 0.15 1
MPR P+ 0.99** 0.2 -0.27 1
MPR P- 0.99** 0.1 -0.17 1
RMD P+ .43** 0.26 0.12 .43** 1
RMD P- .47** 0.12 0.35* .48** 1
GY P+ .65** 0.17 0.53** .64** 0.01 1
GY P- .76** 0.01 0.39* .76** 0.16 1
Screening of Faba Bean (Vicia faba L.) Genotypes for Mycorrhizal Association Parameters
CONCLUSION
Mycorrhizal colonization performance of faba bean genotypes was higher on unfertilized than on P fertilized trial, at both field and greenhouse. Furthermore, the performance of the genotypes was higher at the field than greenhouse condition.
The study indicated that no single genotype was consistently superior for mycorrhizal association across all study conditions. However, Wayu, Dosha, Didea, Moti and Tumsa for phosphorus fertilized; and Dagim, Gebelcho, Dosha, Tumsa and Wayu for the unfertilized trial had performed better. Most of these genotypes were also high yielding. Hence, they may be used for a breeding programme designed to improve mycorrhizal symbiosis and grain yield of the crop. Even though further investigations need to be conducted, the majority of these genotypes are modern genotypes; which is in accordance with previous findings.,
ACKNOWLEDGEMENT
The authors would like to thank Pan African University for funding the research as part of the PhD research project of the first author. Ethiopian Institute of Agricultural Research and Addis Ababa University, under the direct involvement and kind supervision of the third and fourth authors, are duly acknowledged for providing experimental materials, technical support and research spaces.
REFERENCES
Almethyeb M, Ruppel S, Paulsen H-M, Vassilev N, Eichler-Löbermann B (2013). Single and combined
applications of arbuscular mycorrhizal fungi and
Enterobacter radicincitans affect nutrient uptake of faba bean and soil biological characteristics Appl. Agric. Forestry Res.3(63): 229-234
An GH, Kobayashi S, Enoki H, Sonobe K, Muraki M, Karasawa T, Ezawa T (2010). How does arbuscular mycorrhizal colonization vary with host plant genotype?
An example based on maize (Zea mays) germplasms.
Plant Soil. 327:441–453.
Andrade G, De Leij F, Lynch JM (1998). Plant-mediated
interactions between Pseudomonas fluorescens,
Rhizobium leguminosarum and arbuscular mycorrhizae on the pea. Letters in Applied Microbiology 26: 311– 316.
Belay Z, Mauritz Vestberg, Fassil Assefa (2013). Diversity
and abundance of arbuscular mycorrhizal fungi
associated with acacia trees from different land use systems in Ethiopia. Global Journal of Microbiology Research Vol. 1 (1).
Borowicz VA (2001). Do Arbuscular Mycorrhizal Fungi
Alter Plant-Pathogen Relations? Ecol., 82: 3057–
3068.
Brundrett M, Bougher N, Dell B, Grove T, Malajczuk N (1996). Working with Mycorrhizas in Forestry and Agriculture. AClAR Monograph 32. 374 + x p.
Bryla DR, Koide RT (1990). Role of mycorrhizal infection in the growth and reproduction of wild vs. cultivated plants. II. Eight wild accessions and two cultivars of
Lycopersicon esculentum Mill. Oecologia 84:82–92
Cavagnaro TR, Smith FA, Ayling SM, Smith SE (2003). Growth and phosphorus nutrition of a Paris-type arbuscular mycorrhizal symbiosis. New Phytol. 157, 127-134.
CSA (2011). Agricultural sample survey report on farm management practices (private peasant holdings ‘Meher’ season). The FDRE Statistical bulletin, Addis Ababa, Ethiopia.
FAOSTAT (2016). http://www.fao.org/faostat/en/#data/QC Galvan GA, Burger K, Kuyper TW, Kik C, Scholten OE
(2007). Breeding for improved responsiveness to
arbuscular mycorrhizal fungi in onion
Gazey C, Abbott LK, Robson AD (1992). The rate of
development of mycorrhizas affects the onset of
sporulation and production of external hyphae by two species of Acaulospora. Mycol Res 96:643–650 Giovannetti M, Mosse B (1980). An evaluation of
techniques for measuring vesicular arbuscular mycorrhizal infection in roots. New Phytol., 84: 489-500.
Hajiboland R, Aliasgharzad N, Barzeghar R (2009). Influence of arbuscular mycorrhizal fungi on uptake of Zn and P by two contrasting rice genotypes. Plant Soil Environ. 55(3): 93–100
Hildermann I, Messmer M, Dubois D, Boller T, Wiemken
A, Mäder P (2010). Nutrient use efficiency and
arbuscular mycorrhizal root colonisation of winter wheat cultivars in different farming systems of the DOK long‐term trial. J Sci Food Agric 90:2027–2038. Huiying Li, Sally ES, Robert EH, Yongguan Zhu, Andrew
Smith F, (2006). Arbuscular mycorrhizal fungi
contribute to phosphorus uptake by wheat grown in a phosphorus-fixing soil even in the absence of positive growth responses. New Phytologist.172: 536–543 Jia Y, Myles VG, Straker CJ (2004). The Influence of
Rhizobium and Arbuscular Mycorrhizal Fungi on
Nitrogen and Phosphorus Accumulation by Vicia faba. J. Ann. of Bot. 94: 251– 258.
Kaeppler SM, Parke JL, Mueller SM, Senior L, Stuber C, Tracy WF (2000). Variation among maize inbred lines and detection of quantitative trait loci for growth at low
phosphorus and responsiveness to arbuscular
mycorrhizal fungi. Crop Sci 40:358–364
Kucey RMN, Paul EA (1983). Vesicular arbuscular
mycorrhizal spore populations in various
Saskatchewan soils and the effect of inoculation with
Glomus mosseae on faba bean growth in
greenhouse and field trials. Can. J. Soil Sci.63:87-95 Leiser WL, Olotaye MO, Rattunde HF, Neumann G,
Screening of Faba Bean (Vicia faba L.) Genotypes for Mycorrhizal Association Parameters
fungi to improve low-P adaptation of West African sorghums. Plant Soil DOI 10.1007/s11104-015-2437-1 Maalouf F, Ahmed S, Patil S (2017). CH31-Developing
improved varieties of faba bean. Review Meeting.
Maxted, N, Bennett S (2001). Plant genetic resources of legumes in the Mediterranean. Current plant science and biotechnology in agriculture. Kluwer Academic Press, Netherlands. p. 1-386.
Mohamed IAD, Osama NM, Zakaria AMB, Doaa EG (2013). The Promotive effect of Rhizobium
leguminosarum and arbuscular mycorrhizal fungi on
growth and productivity of faba bean plants
cultivated in newly reclaimed sandy soil amended with compost. Journal of Applied Sciences Research, 9(11): 5762-5769, 2013 ISSN 1819- 544.
Mulissa Jida, Fassil Assefa (2012). Phenotypic diversity
and plant growth promoting characteristics of
Mesorhizobium species isolated from faba bean (Vicia faba L.) growing areas of Ethiopia. African Journal of Biotechnology 11: 7483-7493.
Nebiyu A, Jan Diels, Pascal Boeckx (2016). Phosphorus use efficiency of improved faba bean (Vicia faba) varieties in low-input agro-ecosystems. J. Plant Nutr. Soil Sci. 2016, 000, 1–8 DOI:
10.1002/jpln.201500488
Phillips JM, Hayman DS (1970). Improved procedures for clearing roots and staining parasitic and vesicular arbuscular mycorrhizal fungi for rapid assessment of infection. Trans Br MycolSoc 55:158–161
Plenchette C, Fortin JA, Furlan V (1983). Growth responses of several plant species to mycorrhizae in a soil of moderate P-fertility. I. Mycorrhizal dependency under field conditions. Plant and Soil 70:199-209. Saia S, Benitez E, Garcia-Garrido JM, Settanni L, Amato
G, Giambalvo D (2014). The effect of arbuscular mycorrhizal fungi on total plant nitrogen uptake and nitrogen recovery from soil organic material. J. Agric. Sci. 152:370-8.
Sawers RJH, Gebreselassie MN, Janos DP, Paszkowski U (2010). Characterizing variation in mycorrhiza effect among diverse plant varieties. Theor Appl Genet 120:1029–1039. doi:10. 1007/s00122-009-1231-y Schweiger, Peter F, Alan DR, Jim BN, Lyn KA (2007).
Arbuscular mycorrhizal fungi from three genera induce two-phase plant growth responses on a high P-fixing soil. Plant Soil (2007) 292:181–192 DOI 10.1007/s11104-007-9214-8
Singh AK, Hamel C, DePauw RM, Knox RE (2012). Genetic variability in arbuscular mycorrhizal fungi compatibility supports the selection of durum wheat genotypes for enhancing soil ecological services and cropping systems in Canada. Can J Microbiol 58:293– 302
Smith SE, Read DJ (2008): Mycorrhizal Symbiosis. 3rd Edition. Academic Press, London.
Smith SE, Smith FA (2012). Fresh perspectives on the roles of arbuscular mycorrhizal fungi in plant nutrition and growth. Mycologia 104:1–13. doi:10.3852/11-229
Smith SE, Smith FA, Jakobsen I (2003). Mycorrhizal fungi can dominate phosphate supply to plants irrespective of growth responses. Plant Physiol 133:16–20. doi:10.1104/pp. 103. 024380
Tanwar A, Aggarwala A, Panwar V (2013). Arbuscular Mycorrhizal Fungi and Trichoderma viride Mediated Fusarium Wilt Control in Tomato. Biocontrol. Sci. Techn., 23: 485–498.
Tawaraya K (2003) Arbuscular mycorrhizal dependency of different plant species and cultivars. Soil Sci Plant Nutr 49:655–668
Torres AM, Rom B, Avila CM, Satovic Z, Rubiales D, Sillero JC (2006). Faba bean breeding for resistance against biotic stresses: Towards application of marker technology. Euphytica 147:67-80.
van Emden HF, Ball SL, Rao MR (1988). Pest, disease and weed problems in pea, lentil, faba bean and chickpea. In: R.J. Summerfield (Ed.), World Crops: Cool Season Food Legumes, pp. 519–534. Kluwer Academic Publishers, Dordrecht, The Netherlands. Waceke JW, Waudo SW, Sikora R (2001). Response of
Meloidogyne hapla to mycorrhiza fungi inoculation on pyrethrum. Afr. J. Sci. Tech., 2(2): 63-70.
Zhu YG, Smith SE, Barritt AR, Smith FA (2001).
Phosphorus (P) efficiencies and mycorrhizal
responsiveness of old and modern wheat cultivars. Plant Soil 237:249–255.
Zhu, XC, Song FB, Liu SQ, Liu TD, Zhou X (2012). Arbuscular mycorrhizae improve photosynthesis and water status of Zea mays L. under drought stress. Plant Soil Environment.58, 2012 (4): 186–191
Accepted 24 August 2019
Citation: Abu G, Fatokun CA, Keneni G, Assefa F, Belay Z (2019). Screening of Faba Bean (Vicia faba L.) Genotypes for Mycorrhizal Association Parameters. International Journal of Plant Breeding and Crop Science, 6(2): 554-562.
