Field climatic conditions and soil characteristics
The agricultural campaign (2016/2017) was characterized by a severe and long period of drought during the spring season: March, April, May (Fig 1). This drought coincided with the full growth phase of the studied species: durum wheat, when the need for water is high and could therefore affect the nutritional status of the plant and by consequence the association with AMF.
Except for the no-tilled (NT) plot at the Station site, which had a silty texture, the soil in all of the studied plots was clay-silty. EC ranged from 2.23 (ds/m) to 3.42 (ds/m), indicating a slight salinity of the studied soils. The pH ranged from 7.14 to 7.43 indicating a neutral soil in Benifouda and Station and slightly alkaline soil in Remada
(Horneck et al., 2011). The OM content, which ranged between 2.1 and 3.14%, was slightly higher in the NT plots compared to the tilled ones, especially at Remada and Station sites.
The available P amounts (ppm) were low in Remada (8.48) and average in Benifouda (15.74) for both treatments, whereas at the Station site, they were average in the no-tilled plot (15.56) and high in the tilled plot (27.75)
(Horneck et al., 2011). Concerning total N content, the levels recorded in Benifouda (0.23%) were higher than those in Station (0.15%) and Remada (0.16%) sites (Table 2).
AMF root colonization and Spore density
Even if it is not certain that AMF root colonization (RC) is translated into a better performance of the host plant
(Thirkell et al., 2017), high RC rates have often been correlated with increased yields of wheat and other field crops
(Zhang et al., 2019). This suggests that RC could be used to estimate the potential benefits from this symbiosis.
The analysis of variance results (Table 3) showed a significant effect of the site and the tillage system on total colonization rate (TCR), arbuscular colonization rate (ACR) and vesicular colonization rate (VCR), while the combined effect (Site*Tillage) was significant only on TCR and ACR. The mean colonization rates were 31.6, 27.9 and 10.2% for TCR, ACR and VCR, successively. For both Tilled (T) and no-tilled (NT) plots, the TCR recorded in the site “Station” was higher than TCR in “Benifouda” by 59% and TCR in “Remada” by 74%, while the TCR recorded under NT system over all sites was 54% higher than TCR under T system.
Root colonization (RC) depends on the density and affinity of AMF and the host plant, as well as the environmental variables such as soil characteristics and agricultural practices (Gafur, 2014). In our study, RC was positively correlated with organic matter (OM) and pH, and negatively correlated with the available phosphorus amounts (P) in the soil (Table 4). The relation between AMF and OM seems to be synergetic. AMF contribute to the increase of OM in the soil by the AMF biomass generated from the carbon translocated from the host plant, which can reach 20% of the assimilated carbon (Graham, 2000), the microbiome biomass often associated with AMF
(Turrini et al., 2018) and the increase of root biomass as a result of promoting plant growth. On the other hand, organic amendments (compost or manure) stimulate AMF development
(Buto et al., 2016). In contrast to OM, the available P in the soil had a negative correlation with RC. Since it is the main element enhanced by AMF, P excess in soil-root interface or in plant organs can be detected by the host plant, which reacts by inhibiting RC and/or its progress
(Ferrol et al., 2019). While the positive correlation of pH with RC could be explained by the decrease in P availability due to a rise in pH
(Hopkins and Ellsworth, 2005).
Besides soil characteristics, agricultural practices also influence arbuscular mycorrhizal colonization. In general, intensive farming practices have a negative effect on AMF, in contrast to conservation agriculture practices (Säle
et al. 2015). Tillage is important in the intensive crop system with many benefits. However, it can alter the physico-chemical and biological properties of the soil, including AMF
(Siddiqui and Pichtel, 2008). Our findings confirmed the negative impact of tillage on AMF, where the no-tillage improved RC by 54% in durum wheat roots. These results are in accordance with the findings of
Taibi et al., (2020) in similar edaphic and climatic conditions. Tillage affects RC in different ways, as turning over the soil, it buries propagules and/or exposes them to environmental stresses, which will dilute and destroy propagules, and consequently reduce the inoculum potential of the soil. By burying propagules below the depth of early seedling root growth, it delays colonization
(Kabir, 2005). Kabir (2005) also speculated that in NT system, the new formed roots grow following the channels left by the old colonized roots that contains more propagules. The decrease of OM by tillage (Oehl
et al. 2004), could also affect AMF development.
Root colonization of durum wheat in the irrigated site was higher than those in sites under rain-fed conditions. This suggests that water deficit may affect RC as it was reported by Mathur
et al. (2018).
Augé (2001), based on a literature review has concluded that long soil drought decreases RC. Drought can reduce the primary colonization by limiting spore germination
(Wu and Zou, 2017), as it can affect the secondary colonization by inhibiting extra radical hyphal length
(Neumann et al., 2009). In addition, RC could be affected by the carbon supply decrease, because of drought stress on the host plant growth.
Besides their dependence on the AMF species
(Smith and Read, 1997), the storage character of the vesicles suggests that VCR indicates the amount of carbon transferred to AMF by the host plant (Busby
et al. 2012). This can explain the high VCR at the irrigated site, where plants had a better nutritional status than sites exposed to drought.
Moreover, the improvement of RC by no-tillage varied between sites, particularly the arbuscular form (ACR) that ensures nutrients exchange
(Hause and Fester, 2005). This suggests that NT effect depend also on other factors. In the irrigated site (Station), NT improved ACR by 4.76% while in Benifouda and Remada, the improvements were 75.41 and 154.95%, respectively. These results indicate that the NT effect was more important under drought conditions. The increase of water storage in the soil under no-tillage condition
(Huang et al., 2012) could reduce drought impact on RC. In addition, the interesting effect of NT in Remada could be the result of the combined outcome of drought, the tillage practice and the bare fallow (preceding wheat culture), where the removal of host plants reduced arbuscular mycorrhizal establishment
(Schipanski et al., 2014). Bowles et al., (2017) concluded from a meta analysis study that continuous ground cover by crops was as important as the decreasing soil disturbance for AMF.
Spore density (SD), ranged from 3.97 spores/g of soil in the plot under T system in “Benifouda” to 8.2 spores/g soil under T system in “Remada”. Unlike root colonization, tillage effect on SD was not significant despite the decrease by 6,76% (Table 3). NT usually increases spores abundance in soil
(Lehman et al., 2019), which is important particularly for annual crops
(Yang et al., 2014). On the other hand, the site effect was significant on SD (Table 3) with higher density in sites under-fed conditions. This could be explained by the induction of sporulation when soil moisture decreased, which is a specific characteristic allowing AMF adaption to arid conditions
(Jacobson, 1997).