Anatomical characters of faba bean root cells
The root cells of faba bean that grown in narrow or wide ridges before infestation with broomrape contain a cell wall, cortex and vascular cylinder (Fig 1). Before broomrape infestation, the root cells of faba bean allow the passage of water, nutrients and metabolites to the different parts of the plant with no negative impact on the vascular cylinder (transport vessels; xylem and phloem). Meanwhile, broomrape connected to faba bean root by a haustorium that penetrated the epidermal and cortical cells reaching the vascular tissues of faba bean root cells under broomrape infestation. However, this biological situation led to a little negative effect on faba bean root cells as a result of increasing ridge width from 60 to 120 cm, which maintained water, nutrients and metabolites uptake from root cells to different parts of the plant. These results could be due to the wide ridge decreased intra-specific competition between faba bean plants for basic growth resources especially carbon that enhanced physical barriers formation and the cell walls buildup of faba bean root cells, which prevents a haustorium to develop into the root tissues. According to
Hibberd et al., (2002), carbon lost by the parasite through respiration was greater than that of control roots, which explains why broomrape receives large amounts of carbon to grow and develop. In this concern, Joel (2007) revealed that plant haustoria penetrated the host tissues for uptake of organic carbon. It is known that the photochemical apparatus is influenced by high light intensity as mentioned by Critchley (1998) which could reflect on beds than ridges. Consequently, photochemical reactions are linked to the CO
2 fixation process by supplying ATP and NADPH and are also regulated by alternative electron sinks, such as photorespiration, Mehler reaction and nitrogen reduction (Cornic and Fresneau, 2002 and
Noctor et al., 2002). In this concern,
Roth et al., (2005) showed that beds can make plants more energy-consuming because of the increased volume of soil that must be moved in the beds, compared to ridges. Accordingly, there are two mechanisms to tolerate broomrape by increasing ridge width from 60 to 120 cm including the formation of physical barriers against the parasite (Labrousse
et al., 2001) or cell walls build-up and stamping of vascular tissues
(Pérez-De-Luque
et_al2005). These mechanisms could be lead to a low supply of carbon to the parasite through competition with the host plant carbon sinks (Yoder and Scholes, 2010).
Soil total phenols
⮚ Ridge widths
The results in Table 1 show that ridge widths significantly affected soil total phenols concentration in faba bean rhizosphere. Increasing ridge width from 60 to 120 cm increased soil total phenols concentration by 7.32% in the faba bean rhizosphere. The wide ridge may be enhanced efficiency the photosynthetic process of the faba bean plants due to reduction of intra-specific competition between them for basic growth resources compared with those of the narrow one. Additionally, growing faba plants in wide ridges had physical barriers against the parasite and cell walls build-up which probably reflected on a lower supply of carbon to broomrape. Consequently, this biological situation will increase the ability of faba bean plants to consume low amounts of phenols, most of which will be accumulated in the root cells. Conversely, it is likely that intra-specific competition between faba bean plants was increased for basic growth resources than those of the wide one which negatively reflected on photosynthesis and transpiration rates
(Mohamed et al., 2018).
Accordingly, an increase of assimilates in faba bean shoot accompanied with reduced RNase and protease activity, especially polyphenols will be bonded with other organic substances such as carbohydrate or protein
(Saharan et al., 2002). Seven phenolic acids were detected in the rhizosphere that continuously cropped faba bean
(Dong et al., 2016) where faba bean seeds contain several other bioactive compounds, such as polyphenols
(Turco et al., 2016).
⮚ Cropping systems
Soil total phenols concentration in faba bean rhizosphere was significantly affected by the cropping systems (Table 1). 50% faba bean + 100% fenugreek recorded higher soil total phenols concentration than the others, meanwhile, the converse was true for 100% sole faba bean (Table 1). No doubt, increasing fenugreek plant density from 50 to 100% of sole cropping increased total phenols concentration in faba bean rhizosphere. It is important to mention that higher plants produce many phytochemical constituents that not only play a pivotal role in plants producing them but also affect their neighboring plant communities (Madany and Khalil, 2017). Also, they reported that the allelochemical components of fenugreek seed extract were nine phenolic compounds.
Accordingly, fenugreek roots secreted large quantities of total phenols in faba bean rhizosphere due to low consumption of total phenols in their metabolic reactions, meanwhile, faba bean roots secreted large quantities of total phenols in their rhizosphere (Table 1). So, it may be possible that soil total phenols concentration was increased by increasing fenugreek plant density with decreasing faba bean plant density per unit area which explains increased soil total phenols concentration with 100% sole faba bean. These results are confirmed by
Candido et al., (2016) who found that allelochemicals
(i.e. phenolic and terpenoid constituents) released into the environment either by exudation from roots or leaching from the aerial parts and affect the structure of the plant community.
⮚ The interaction between ridge widths and cropping systems
Soil total phenols concentration in faba bean rhizosphere was significantly affected by the interaction between ridge widths and cropping systems (Table 1). Growing 50% faba bean + 100% fenugreek in wide ridges had higher soil total phenols concentration than the others in faba bean rhizosphere, meanwhile, the reverse was true for growing 100% sole faba bean in the narrow ones.
Broomrape infestation and leaf chlorophylls a and b
⮚ Ridge widths
The results in Table 2 show that ridge widths significantly affected number of spikes per m
2 and spikes dry weight per m
2 and leaf chlorophylls a and b in both seasons. Expressing tolerance by number of spikes per m
2 and dry weight of spikes per m
2 showed considerable variation between two ridge widths. Growing faba bean in wide ridges reduced number of spikes per m
2 and dry weight of spikes per m
2 by 15.90 and 25.84% in the first season and 19.14 and 28.11% in the second one, respectively, compared with those grown in the narrow ones. Meanwhile, increasing ridge width from 60 to 120 cm increased leaf chlorophylls a and b by 4.78 and 7.81% in the first season and 4.05 and 6.77% in the second one, respectively, under heavily soil infested with broomrape. These results show that the wide ridges played a major role in maintaining soil moisture which provided suitable conditions for early germination and growth of faba bean seedlings compared with those grown in the narrow ones, indicating escaping from severe infestation with broomrape. A flat-topped ridge, similar to a wide bed, conserved moisture better than a sharply sloped ridge and hastened emergence by four to five days (Prestt and Carr, 1984). Also, they showed that dry areas of sharply sloped ridges tended to form beneath the sides of the ridges and the steep sides encouraged runoff. This means that more irrigation water probably flowed into the depths of the soil by growing faba bean in the narrow ridge compared with those grown in the wide one, which induces broomrape seeds for germination. Moreover, growing faba bean plants in wide ridges had physical barriers with stamping of the vascular system in the root cells that probably reflected on a lower supply of carbon to broomrape (Fig 1). It is known that the seeds need to be exposed to a moist environment (called preconditioning) for several days at a suitable temperature (optimum 15 to 20°C) before seeds respond to germination stimulants (Kebreab and Murdoch, 1999). These results are in parallel with Safina (2017) who showed that there was more shading around the plants of the narrowest ridge (moist environment) than those of the widest ridge. Also, Zohry and Ouda (2017) reported that germination and growth of faba bean seedlings of the widest ridge were faster than those grown of the narrowest one. In the same trend, increasing wide ridge width from 60 to 120 cm increased total phenols concentration in the rhizosphere of faba bean roots (Table 1) which reduced broomrape growth and maintained leaf chlorophylls pigment from deterioration. These results are in parallel with
Mohamed et al., (2018) who showed that faba bean plants of the wide ridge had higher shoot chlorophylls a and b concentrations than those of the narrow one.
⮚Cropping systems
The results in Table 2 show that cropping systems significantly affected number of spikes per m
2 and spikes dry weight per m
2 and leaf chlorophylls a and b in both seasons. All intercropping treatments of fenugreek with faba bean had a lower number of spikes per m
2 and spikes dry weight per m
2 and higher leaf chlorophylls a and b than sole faba bean in both seasons. In other words, 50% faba bean + 100% fenugreek had lower number of spikes per m
2 and spikes dry weight per m
2 by 69.42 and 68.49% in the first season and 51.73 and 51.88% in the second one, respectively, compared with 50% faba bean + 50% fenugreek. Also, 50% faba bean + 100% fenugreek increased leaf chlorophylls a and b by 2.63 and 20.00% in the first season and 2.69 and 20.31% in the second season, respectively, compared with 50% faba bean + 50% fenugreek. Meanwhile, 100% faba bean + 100% fenugreek decreased number of spikes per m
2 and spikes dry weight per m
2 by 33.39 and 18.94 % in the first season and 45.94 and 31.73 % in the second season, respectively, compared with 100% faba bean + 50% fenugreek. Meanwhile, 100% faba bean + 100% fenugreek increased leaf chlorophylls a and b by 2.20 and 15.15% in the first season and 2.28 and 14.51% in the second season, respectively, compared with 100% faba bean + 50% fenugreek. These results were attributed to increasing fenugreek plant density per unit area from 50 to 100% of sole cropping in the two cropping systems (50% faba bean + 100% fenugreek and 100% faba bean + 100% fenugreek) increased soil total phenols concentration in faba bean rhizosphere (Table 1). Moreover, decreasing faba bean plant density from 100 to 50% of sole cropping could be reduced stimulators of broomrape seed germination by increasing soil total phenols concentration in faba bean rhizosphere due to reduce intra-specific competition between faba bean plants for basic growth resources. Particularly,
Bouwmeester et al., (2003) and
Plakhine et al., (2009) indicated that the germination of these obligate root parasites depends on the receipt of a chemical signal, which is present in the rhizosphere of potential hosts. So, increasing fenugreek plant density increased soil total phenols concentration in faba bean rhizosphere which negatively affected broomrape seed germination under heavily soil infested with broomrape. Also, it is expected that the broomrape that penetrated faba bean root cells, its life cycle will not be completed as a result of the accumulation of the total phenols in faba bean root cells. Inhibition of
Orobanche crenata seed germination by allelochemicals released by fenugreek roots is the mechanism for reduction of this parasite
(Fernández-Aparicio
et_al2008 and 2011). This biological situation will maintain leaf chlorophyll pigment from deterioration. It is known that the total leaf area of faba bean seedlings was significantly increased by increasing fenugreek extract concentration (Madany and Khalil, 2017), meanwhile, high densities of faba bean plants can cause the plants to senesce earlier (Gezahegn, 2019).
⮚ The interaction between ridge widths and cropping systems
Number of spikes per m
2 and their dry weight per m
2 and leaf chlorophyll b were significantly affected by the interaction between ridge widths and cropping systems in both seasons (Table 2). Growing 50% faba bean + 100% fenugreek in wide ridges had a lower number of spikes per m
2 and spikes dry weight per m
2 and a higher leaf chlorophyll b followed by 100% faba bean + 100% fenugreek, meanwhile, the converse was true for growing 100% sole faba bean in the narrow one. These results were attributed to formation of physical barriers with stamping of the vascular system in the root cells (Fig 1) and higher total phenols in faba bean rhizosphere (Table 1) which reduced broomrape growth and maintained leaf chlorophyll pigment from deterioration (Table 2). This interaction probably reflected on a lower supply of carbon to the parasite.
Faba bean seed yield and its attributes
⮚Ridge widths
Ridge widths significantly affected the percentage of the harvested plants, plant height, seed yield per plant, 100-seed weight and seed yield per ha in both seasons, meanwhile numbers of branches and pods per plant were not affected (Table 3). Increasing ridge width from 60 to 120 cm increased the percentage of the harvested plants, seed yield per plant, 100-seed weight and seed yield per ha than the other one, meanwhile, the converse was true for plant height. In other words, growing faba bean in wide ridges gave a higher percentage of the harvested plants by 12.45 and 11.46% in the first and second seasons, respectively, than the other one. Also, growing faba bean in wide ridges had a higher seed yield per plant by 16.18 and 26.20% in the first and second seasons, respectively, than the other one. Moreover, growing faba bean in wide ridges had a higher 100-seed weight by 4.19 and 3.12% in the first and second seasons, respectively, than the other one. Furthermore, growing faba bean in wide ridges had a higher seed yield per ha by 19.48 and 20.73% in the first and second seasons, respectively, than the other one. These results were due to lower infestation with broomrape by growing faba bean plants of the wide ridges that had a higher ability to maintain soil moisture which furnished suitable environmental conditions for germination and early growth of faba bean seedlings than growing faba bean in the narrow one. In this concern, Zohry and Ouda (2017) showed that the widest ridge had high moisture which provided suitable conditions for germination and early growth of faba bean seedlings compared to those grown in the narrowest one, thus escaping from severe infestation with broomrape. Also, growing faba bean in wide ridges had physical barriers with stamping of the vascular system in the faba bean root cells (Fig 1) which reduced broomrape growth and maintained leaf chlorophyll pigment from deterioration (Table 2). These defense mechanisms lead to high dry matter accumulation in different tissues of the plant during growth and development. These results are in the same context as
Fisher et al., (1995) who found increased yield in wide beds attributable to moisture conservation in the beds. Also, Safina (2017) reported that increasing ridge width from 60 to 120 cm increased the number of pods per plant, seed yield per plant, seed index and seed yield per ha by 5.68, 3.02, 4.98 and 5.20%, respectively. Moreover,
Mohamed et al., (2018) reported that growing faba bean in wide ridges had lower values of plant height and higher values of seed yields per plant and per ha than those of the narrow one.
⮚ Cropping systems
Cropping systems significantly affected the percentage of the harvested plants, plant height, seed yield per plant, 100-seed weight and seed yield per ha in both seasons, meanwhile numbers of branches and pods per plant were not affected (Table 3). A lower percentage of the harvested plants and seed yield per ha was achieved by 50% sole faba bean than the others. These results were due to a lower soil total phenols concentration in faba bean rhizosphere (Table 1) which led to higher infestation with broomrape and maintained leaf chlorophyll pigment from deterioration (Table 2).
Orobanche induced growth inhibition of faba bean due to parasitism might result from depletion of host nutrition (Nemat
Alla et al., 2007). Orobanche induced yield reductions are not primarily due to competition for water, but rather due to carbohydrate loss to the parasite
(Bayoumi et al., 2014). Conversely, 100% faba bean + 100% fenugreek had a higher percentage of the harvested plants, plant height and seed yield per ha than the others. These results were attributed to increase faba bean plant density from 50 to 100% of sole cropping increased seed yield per hectare. This biological situation increased plant height. Meanwhile, 50% faba bean + 100% fenugreek had a higher seed yield per plant and 100-seed weight than the others. These results were due to 50% faba bean + 100% fenugreek increased soil total phenols concentration in faba bean rhizosphere (Table 1) which reduced infestation with broomrape and increased leaf chlorophylls a and b during growth and development (Table 2). These results are in harmony with
Elsakhawy et al., (2020) who showed that plant height, pod number per plant, seed yield per plant and 100-seed weight were significantly decreased under broomrape infestation.
⮚The interaction between ridge widths and cropping systems
The interaction between ridge widths and cropping systems significantly affected seed yield per plant in the first season, meanwhile, the other studied traits were not affected (Table 3). Growing 50% faba bean + 100% fenugreek in wide ridges had a higher seed yield per plant than the others. These results were due to wide ridges furnished suitable environmental conditions for tolerance of broomrape infestation (Table 1). This tolerance was improved by increasing fenugreek plant density from 50 to 100% of sole cropping which reduced number of spikes per m
2 and spikes dry weight per m
2 and increased leaf chlorophylls a and b (Table 2).
Fenugreek seed yield and its attributes
⮚ Ridge widths
Ridge widths significantly affected plant height, seed yields per plant and per ha in both seasons, meanwhile numbers of branches and pods per plant were not affected (Table 4). Increasing ridge width from 60 to 120 cm increased seed yields per plant and per ha in both seasons, meanwhile, the converse was true for plant height. Growing fenugreek in wide ridges gave higher seed yields per plant and per ha by 17.26 and 15.69% in the first season and 16.75 and 17.10% in the second season, respectively, than those grown in the narrow one. These results may be due to wide ridges provided suitable environmental conditions for germination and early growth of fenugreek seedlings compared with those grown in the narrow one. These results are in parallel with Tuncturk (2011) who showed that some yield parameters were increased with increasing row spacing.
⮚Cropping systems
Cropping systems significantly affected plant height, seed yields per plant and per ha in both seasons, meanwhile numbers of branches and pods per plant were not affected (Table 4). Higher plant height and seed yield per plant was obtained by 50% of sole fenugreek than the others. These results could be due to lower intra-specific competition between fenugreek plants for basic growth resources which positively reflected on dry matter accumulation during growth and development. However, increasing fenugreek plant density from 50 to 100% of sole cropping did not increase number of pods per plant. These results are in accordance with Bismillah
Khan et al., (2005) who showed that spatial arrangement did not have any significant effect on number of pods per plant. Meanwhile, 100% faba bean + 100% fenugreek had higher plant height and lower seed yield per plant than the others. These results could be due to increase fenugreek plant density from 50 to 100% of sole cropping increased inter and intra-specific competition between both and same species, respectively, for basic growth resources which negatively affected photosynthesis process. However, 100% faba bean + 50% fenugreek had a lower seed yield per ha than the others. These results may be attributed to increase faba bean plant density from 50 to 100% of sole cropping had negative effects on metabolic reactions in different tissues of fenugreek plant, this biological situation positively interacted with lower fenugreek plant density per unit area to reduce photosynthesis process and dry matter accumulation during growth and development.
⮚ The interaction between ridge widths and cropping systems
The interaction between ridge widths and cropping systems significantly affected seed yield per plant in the first season, meanwhile, the other studied traits were not affected (Table 4). Growing 50% or 100% sole fenugreek in wide ridges recorded a higher seed yield per plant than the others. These results demonstrate that wide ridges decreased intra-specific competition between fenugreek plants for basic growth resources, which increased dry matter accumulation during growth and development. These results are confirmed with
Tiwari et al., (2016) who indicated that significantly higher seed yield (494.45 kg/ ha) was observed with a plant spacing of 20 cm ´ 10 cm than the others.
Competitive relationships and intercropping economic advantage
⮚Land equivalent ratio (LER)
The total LER values were better than one in all the studied treatments. Fenugreek + faba bean in narrow or wide ridges increased LER compared with sole plantings of faba bean (Table 5). LER ranged from 2.63 by growing 50% fenugreek + 50% faba bean in narrow ridges to 4.39 by growing 100% fenugreek + 100% faba bean in wide ridges in the first season. Also, LER ranged from 1.93 by growing 50% fenugreek + 50% faba bean in narrow ridges to 3.33 by growing 100% fenugreek + 100% faba bean in wide ridges in the second season. The advantage of higher LER by fenugreek + faba bean in wide ridges over sole plantings of faba bean were due to wide ridge formed physical barriers with stamping of the vascular system in the root cells (Fig 1) and higher soil total phenols concentration in faba bean rhizosphere (Table 1) which decreased infestation with broomrape and maintained leaf chlorophyll pigment from deterioration (Table 2). This biological situation reduced intra-specific competition between faba bean plants and broomrape for basic growth resources. These results are in accordance with Safina (2017) who found that LER ranged from 1.63 for flax + Giza-2 variety in ridges 60 cm width.
⮚ Intercropping economic advantage
Data presented in Table (5) indicate that fenugreek + faba bean in narrow or wide ridges was more profitable to farmers than sole plantings of faba bean in both seasons. Total return varied from 1484.76 USD/ha by growing 50% fenugreek + 50% faba bean in narrow ridges to 2478.52 USD/ha by growing 100% fenugreek + 100% faba bean in wide ridges in the first season. Also, total return varied from 1057.18 USD/ha by growing 50% fenugreek + 50% faba bean in narrow ridges to 1826.57 USD/ha by growing 100% fenugreek + 100% faba bean in wide ridges in the second season. With respect to MAI, it varied from 920.21 by growing 50% fenugreek + 50% faba bean in narrow ridges to 1913.93 by growing 100% fenugreek + 100% faba bean in wide ridges in the first season Also, MAI varied from 509.41 by growing 50% fenugreek + 50% faba bean in narrow ridges to 1278.05 by growing 100% fenugreek + 100% faba bean in wide ridges in the second season. It is important to mention that 100% fenugreek + 100% faba bean with wide ridges recorded the highest economic return compared with the others, indicating this system was more profitable to the farmers than the others and it should be recommended under heavily soil infested with broomrape.