Resistance to Orobanche crenata Forsk. in lentil (Lens culinaris Medik.): exploring some potential altered physiological and biochemical defense mechanisms

ABSTRACT Management of broomrape (Orobanche crenata Fosk.) that causes important damages on lentil production becomes a veritable concern in the Mediterranean region. Eighty lentil accessions were evaluated for resistance to O. crenata under field and controlled conditions. Both genotypes ILL6415 and ILL7723 expressed the highest resistance level under field and pot experiment with low Orobanche infestation and relatively high seed yield (50.1 g m−2). Such resistance was associated with physiological and biochemical changes in metabolites profiling. In total, 109 and 115 metabolites were identified in the lipophilic phase of both ILL6415 and ILL7723, respectively, against only 92 metabolites recorded for susceptible check Zaaria. Significant differences were observed in metabolite concentrations (fatty acids, sterols alkanes) between roots and shoots of susceptible and resistant infested plants. Accumulation of α-linolenic acid and arachidic acid was more pronounced in the resistant genotypes ILL6415, ILL7723 which could be associated with resistance pathways involved in the resistance to O. crenata. GRAPHICAL ABSTRACT


Introduction
Lentil (Lens culinaris Medik.) is the third-most important cool-season grain legume in the world after chickpea and pea. It is widely grown as a rainfed crop on 3.85 million hectares area and accounted for 6% of the total global pulses production with 3.59 million tonnes and average yield of 0.93 tn ha −1 (Sehgal et al. 2021). The major production regions are South Asia and China (44.3%), North America (41%), Central and West Asia and North Africa -CWANA (6.7%), Sub-Saharan Africa (3.5%) and Australia (2.5%) (Kumar et al. 2013). Lentil has been cultivated for more than 10,000 years for its important agronomic and socioeconomic roles worldwide (Erskine et al. 2011). With high protein level and an important amount of carbohydrates, fibers, minerals and antioxidant compounds, lentil is considered as one of the most nutritious legume crop (Singh and Singh 2014). It is an important staple food crop particularly among the poor populations and smallholder farmers (Çarman 1996). As other legumes crops, lentil is considered as a pivotal component for sustainable agriculture due to its ability to fix nitrogen in the soil, which makes practical to use it in rotation with cereals (Shah et al. 2003). Morocco is the second major producer of lentil in Africa with more than 40,000 ha cultivated area (Taha et al. 2018), and total annual production of 30,670 tons (Idrissi et al. 2020). It account about 14% of the total legume cultivated area in the country. In Morocco, Lentil is mainly cultivated under rainfed conditions in the dry areas where drought is the major abiotic constraints limiting the crop production and productivity due to low and irregular rainfall (Idrissi et al. 2020). Attack by the parasitic weed O. crenata seems to be the most important biotic stress limiting the production and the development of the crop especialy in Zaair region which is the most important production area in the country. In fact, O. crenata is an holoparasitic plants completely dependent on the host for its nutritional requirements. Its is considered as a serious threat that causes important damages and yield losses on many legume crops in the Mediterranean region and Sub-Sahan Africa (Amri et al. 2021). During the last decades, attack by this parasite has burden lentil production and productivity in Morocco (Kumar et al. 2015;Abu-Irmaileh and Labrada 2017;Idrissi et al. 2020). The continuous spread of this threat and important damages that causes on lentil and other host crops often force farmers to give up growing these crops . Recent assessment conducted in Morocco reported 51% estimated infested legumes cultivated areas with an average yield loss of 30-40% (Abu-Irmaileh and Labrada 2017).
Several control methods have been tested but none of them resulted in complete and successful control of the parasite . Breeding for resistance is considered as the most economically feasible and environment friendly control method . This strategy has shown promising success and many resistance sources were identified in faba bean, Chickpea, sunflower and tomato (Kharrat et al. 2010;Nefzi et al. 2016;Amri et al. 2019;Bai et al. 2020;Cvejić et al. 2020). However, breeding for resistance to broomrape remains complicated because of the limited source of the resistance and the low heritability of the genes and QTLs associated with that resistance (Pérez- De-Luque et al. 2005;Amri et al. 2021). Indeed, a better understanding and development of solid knowledge about the resistance mechanisms involved and the interaction between the host and the parasite will help to improve the resistance level and develop new breeding material (Pérez-De-Luque et al. 2007;Trabelsi et al. 2017;Abbes et al. 2020). Lentil is poorly competitive with O. crenata, which can cause complete yield loss under high infested conditions. Unfortunately, studies on lentil resistance to O. crenata and the interaction between host and parasite are still insufficient. A recent scientific report showed that the metabolomics profile of the parasite is different from that of the host, which means that the parasite may have a self-regulating in metabolism (Amir 2016; Clermont et al. 2019). Some other studies reported that environmental stresses cause changes in primary and secondary plant metabolisms which depend on plant resistance strategies (Hasanuzzaman et al. 2013).
In this study, we aimed to evaluate the response of a lentil germplasm collection to O. crenata parasitism under field and controlled conditions and investigate the potential metabolic differences between identified susceptible and resistant genotypes.

Plant material and field trials
In total, 80 lentil genotypes were subjected to field evaluation and screening for resistance to O. crenata (Table 1). Germplasm was provided by the International Center of Agricultural Research in dry areas (ICARDA) genebank. The trial was conducted during two consecutive cropping seasons 2017/2019 in a high O. crenata infested sick plot at ICARDA Merchouch research station -Morocco. Different genotypes were planted end of November according to an alpha lattice design with two replications. For each genotype, 30 seeds were planted in 1 m row with 30 cm inter-row spacing. The local cv. Bakria which is reported to be moderately susceptible to O. crenata was used as check. Hand weeding was done when necessary and neither herbicide nor fertilizer were applied. The following parameters were recorded before and at crop maturity, Number of days to flowering (D2F), days to Orobanche emergence (D2OE), Orobanche incidence (OIN), Orobanche severity (OSV), Emerged Orobanche number per plant (EON), Emerged Orobanche dry weight per plant (EODW), biological yield g m −2 (BY) and seed yield g m −2 (SY), the harvest index (HI) and parasitism index (PI). The PI was calculated according to the following formula: OIN: Percentage of lentil plants showing at least one emerged shoots of orobanche per row. OSV: level of damage (1-9 scale) caused by the parasite on lentil growth and seed production (Abbes et al. 2007).

Pot experiment
Out of the 80 tested genotypes, five lines selected for their resistance to O. crenata were subjected to a confirmation pot experiment under controlled conditions with two Orobanche treatments (infested and non-infested). The five selected lines all with two released varieties (Bakria and Zaaria) were planted in 2 l pots. Infested pots were inoculated with O. crenata seeds (collected during previous seasons on faba bean plants) at a density of 20 mg.kg −1 soil. Four replications were considered for each genotype/treatment and planting was performed mid-November with 3-4 seeds per pot. Pots were watered when necessary to keep plants at good soil moisture. After emergence, the number of lentil plants was reduced to only one plant per pot. At the end of the experiment, the 56 days aged plants were uprooted from pots. The host root system all with O. crenata attachments were washed carefully and the following parameters were determined. O. crenata attachments were counted and classified according to their development stage (Abbes et al. 2011) to underground/non-emerged Orobanche tubercles (NEO) and emerged Orobanche shoots (EON) per plant. Lentil shoot (SDW) and root (RDW) dry weight were also determined for the same plants.

Metabolites analyses
Untargeted metabolomics profiling was performed using gas chromatography-mass spectrometry (GC-MS). The method used in this study for organic extraction and transesterification from shoots and roots of both infested and non-infested plant is sited by (Mutale-joan et al. 2020) with some modifications. Samples were grinded in liquid nitrogen then 300 mg from each sample dissolved in 4 mL chloroform/ methanol mixture (2/1, v/v). 10 µL of internal standard Dodecane (Sigmaaldrich; Density, 0.75 g/mL) were added into the mixture, which were placed to a heat block (Labet International, Edison, USA) pre-set at 85°C and left for 2 h. After the mixture was placed to an ultra-sound bath (Branson ultrasonic Sonifier 450, Danbury, USA), the sonication was carried for 60 min. One milliliter of H 2 O was added to the vials and the mixtures were thoroughly vortexed. Then the organic phase was transferred into a new vial and the chloroform was evaporated under a stream of nitrogen gas. For acid transesterification, 500 µL of methanol/sulfuric acid (6%, v/v) was added to dried organic material, then the mixture was heated, sonicated and dried as described above. 750 µL of chloroform and 250 µL of distilled water were added for phase separation. The organic phase was collected and conserved at −20°C for further analysis. The gas chromatography (GC) (Agilent 7890A Series) coupled to mass spectrometry (MS) was used for the identification of apolaire compounds. A volume of 4 μL of each tested samples were injected into the 123-BD11 column (15 m × 320 μm × 0.1 μm) by 1/4 split mode using helium as carrier gas at 3 mL.min −1 . Briefly, the temperature was set at 230 and 150°C in the ion source and the MS transfert line, respectively. The oven temperature was set to started at 30°C and to reach 360°C at the end. The identification of metabolite was carried by the comparison of their mass spectra (MS) with NIST 2014 MS Library.

Statistical analysis
Statistical analyses were done using SPSS software and R script. Analysis of variance (ANOVA) for field data was done considering a linear mixed model with blocks and repetitions as random factors and seasons and genotypes as fixed factors. Person correlation coefficients between all morphological traits were computed and tested for their statistical significance using prcomp function and Mix-Omics package. PCA and heatmap was generated using R studio, visualization of corrplot and ggplot packages, integrated into the R sofware. One way anova and Duncan's test was studied using SPSS software.

Field screening for resistance to O. crenata in lentil
Results showed a high variability within the lentil germplasm collection in the response to O. crenata parasitism. Field screening and evaluation showed significant differences (P ≤ 0.05) between the tested genotypes for D2F, D2OE, PI, BY, SY and HI. No significant differences were recorded for EON and EODW. The cropping season showed significant effect on D2F, D2OE, EON, EODW, SY and HI and no significant effect on PI and BY. Except for PI, the interaction genotype*cropping season was significant for all other parameters ( Table 2). Out of the 80 tested accessions, only five accessions showed a good resistance level to O. crenata. The correlation matrix ( Figure 1) shows that traits related to lentil productivity such as SY and HI were negatively correlated with traits related to O. crenata infestation level D2OE (r = −0.20**), PI (r = −0.37**) and EON (r = −0.34**). Interestingly negative correlations were found also between SY and HI (r = −0.42**) and D2F (r = −0.56**) ( Figure 1). However, a high positive correlation was found between D2F and D2OE, D2OE and EODW, SY and BY, BY and D2OE, EON and PI (Correlation coefficients varied between 0.17 and 0.68, p ≤ 0.001). The correlations between different traits were confirmed at a higher dimension using PCA which was used to select the best resistant genotypes. Five genotypes (ILL6415, ILL1861, LIRL21187, ILL7723, and ILL4830) were identified with a good level of resistance to O. crenata under field conditions. PCA revealed that the first three principal components (PC) explained 81% of the total original variation. The first PC, which explained 38.8% of the total variation, showed negative correlation with PI (r = −0.63) and positive correlation with BY (r = 0.57), SY (r = 0.89) and HI (r = 0.85). This PC corresponds to the list of genotypes with, high PI and low BY and SY such as ILL5645 and ILL82. The second PC, which described 24.6% of the total variability, is characterized with a positive correlation with EON (r = 0.76) and EODW (r = 0.76) and opposed the genotypes with high EON and EODW such as ILL5418 and ILL705 to those with low number such as ILL6415 and ILL7723. The third PC explained 17.6% of the total data variability and it is positively correlated with D2F (r = 0.62) and EODW (r = 0.64) which corresponds to genotypes with high D2F and D2OE ( Figure  2).
A cluster analysis, using all collected data was performed to cluster the different studied genotypes based on their resistance level. Four groups were identified ( Figure 2). The first cluster contains 46 genotypes that presented a high susceptibility to O. crenata with high PI values ranging from 5.53 to 9 and low HI and seed production levels (SY ≤ 36.5 g m −2 ). The second cluster with only six genotypes, showed a moderate susceptibility to O. crenata expressed by high D2OE, a PI varying from 6.9 to 8.75, moderate emerged Orobanche number and dry weight. The level of seed production for the genotypes in this cluster varied from 6.22 to 39.1 g m −2 with a maximum HI of 0.   with the highest resistance level were grouped in the fourth cluster. This group was characterized by relatively the lowest PI ranging from 2.77 and 5.87. The best recorded SY were observed for the genotypes of this cluster such as ILL7723, ILL4615, ILL1861and LIRL21187. Genotypes from cluster four showed the best resistance level against O. crenata infectation and relatively high seed production (Table 3).

Pot experiment and confirmation of the resistance under controlled conditions
Five genotypes from cluster 4, ILL1861, ILL4830, ILL4615, ILL7723 and LIRL21187, were selected to be subjected to a confirmation experiment and evaluation of the impact of the parasite on host development under controlled conditions. Result showed a highly significant difference between the genotypes (p < 0.001) under both infested and free Orobanche conditions. Compared to non-infested control plants, biomass production (shoots and roots) was significantly decreased by O. crenata for all the tested genotypes except ILL6415 (Figure 3(A)). However, no significant differences were observed between the studied genotypes for the average number of TON (p > 0.05) that varied from 3.5 to 7.2, and the NEO (p > 0.05) wish varied from 3.2 to 5.9. Only EON showed significant differences between genotypes (p ≤ 0.05) wish varied from 0.4 to 0.7 (Table 4).
Significant differences were observed between tested genotypes under both treatments (infested and non-infested) (p < 0.001) for the SDW (Table 5). Also, results showed that O. crenata significantly affected the RDW (p ≤ 0.05). The genotypes*treatment interaction was highly significant for SDW (p < 0.001) and not significant for RDW (p > 0.05). Compared to non infested plants, an important decrease of SDW (90%) was observed for genotypes Bakria, Zaaria and ILL4830. Moderate to low SDW decreases were observed for the genotypes ILL6415 (53.8%) and ILL7723 (66%). The same genotypes showed a relatively low RDW decrease with respectively 19.2% and 22.2% against a maximum of 72% recorded for Zaaria and ILL4830.

Qualitative untargeted metabolomic analysis
To investigate the impact of O. crenata parasitism on the metabolomic profile of susceptible and resistant lentil genotypes and to gain more information and knowledge about the chemical and biochemical mechanisms involved in the resistance mechanisms, a metabolic profiles analysis was performed using GC-MS for three lentil genotypes characterized by different resistance levels; two resistant genotypes (ILL6415, ILL 7723) and one susceptible genotype The metabolites concentration found in the shoot and the root of infested and non-infested genotypes were used to perform a PCA and to build heat-maps to identify the pattern between the samples and gain clear details about the metabolic changes (Figures 4 and 5). The PC1 and PC2 of the shoot metabolites explain 78% and 17.6% of the total variance respectively. For both genotypes ILL6415 and ILL7723, the non-infested and infested pant functions were close to each other. Instead, PC1 and PC2 separate the function of the non-infested and infested Zaaria (Figure 4(B)). Differently to the shoot, PC1 and PC 2 explain 36.3% and 25.3% of the total root metabolite variations. The PC plot separates markedly the functions of non-infested root of ILL6415, ILL7723 and Zaaria from the infested root functions ( Figure  5(B)). Out of 19 abundant and identified metabolites detected, only linolenic acid, arachidic acid and abietic acid are markedly accumulated in the shoot of the resistant infested genotypes ILL6415. Four metabolite levels including stigmasterol, methyl 2-hydroxytetracosanoate, lignoceric acid and stigmastan-3,5-diene decreased compared with the non infested plants. On the other hand, the metabolite concentrations do not show significant changes between the treatments in the shoot of the resistant genotype ILL7723. While linolenic acid, g-sitosterol and melissic acid exhibit increased concentrations in the shoot of the infested susceptible genotypes Zaaria compared with the noninfested plants (Figure 4(A)). Moreover, concentrations of g-sitosterol, stigmastan-3,5diene, telfairic acid, docosanoic acid, stigmasterol and arachidic acid significantly increased in the infested root of ILL6415, when a significant reduction of palmitic acid, oleic acid, lignoceric acid and methyl 2-hydroxy-tetracosanoate were found in the same infested roots. Stearic acid, telfairic acid and montanic acid were more abundant in the infested root of ILL7723 than the non-infested root, for the same genotype only melissic acid and palmitoloic acid were reduced in the infested root compared to the noninfested root. For the susceptible genotype Zaaria, tricosanoic acid and pentacosanoic acid were significantly accumulated in the infested root ( Figure 5(A)).

Field evaluation
The holoparasitic weed O.crenata is known for its devastating effect on productivity of many crop species, especially legume crops such as faba bean, chickpea, grass pea and lentil   (Ennami et al. 2017;Mbasani-Mansi et al. 2019). Regarding our results, early flowering was positively correlated to D2OE (r = 0.68***) and negatively correlated with SY (r = −0.20***). The early flowering, which has been described to have a positive effect on plant potential   A and B). Metabolite concentrations were row normalized to highlight differences among treatments. Relative abundance ranges from blue (lower than the average percentage value) to red (higher than the average percentage value). The Compounds identified in the different apolar extracts derived from infested and non-infested shoot were grouped in structurally related families (C).
yield through the prolongation of reproductive phase as well as seed filling period (Sundaram et al. 2019), is associated with genes that are highly affected by the environmental conditions (Yaish et al. 2011). It is tightly controlled by some complex genetic and metabolic pathways such as salicylic acid and gibberellin pathways as well as some other phytohormones involved in plant growth (Dezar et al. 2011;Jing et al. 2020). The root parasitic plants are metabolically dependent on their hosts (Clermont et al. 2019), which suggest that the early metabolism adjustments associated with early flowering will stimulate the release of germination stimulants in the rhizosphere of the host root system resulting in an early attachment and development of the parasite on the host plant. such early infestation results with an early parasitism impact affecting the host metabolism and thus its physiological and phenological behavior which explain the correlation found in the current study between D2F, D2OE and SY (Amri et al. 2021). Parasitism Index was positively correlated with EON and EODW, which reflects the severe negative effect of O. crenata parasitism on lentil growth and seed production. Similar results were reported in previous studies performed on faba bean (Abbes et al. 2011;Trabelsi et al. 2015;Trabelsi et al. 2016), lentil (Ennami et al. 2017), chickpea (Nefzi et al. 2016) and grass pea (Abdallah et al. 2020).
The score plot, the biplot and cluster analysis are graphical representations used for a better visualization of PCA results, and allow the identification of genotypes with the desirable characteristics. In general, whatever the genotypes are located close to the trait that's mean a higher value for the corresponding trait. The genotypes ILL4605, ILL4830, LIR22107, ILL6415 and ILL7723 clustered together in cluster 4 showed the lowest PI levels and the highest SY and BY. The same genotypes are located to the negative side of PC2 which means they have low values of EON and EODW. On the other hand, genotypes such as ILL82, ILL304, ILL5385 clustered together in cluster 1, showed the highest PI and lowest SY and BY, and presented as the most susceptible genotypes to O. crenata. Figure 5. Heat map and principal component analysis (PCA) of metabolite levels detected in infested and non-infested lentil genotypes (centered and scaled) of root metabolites. Metabolite concentrations were row normalized to highlight differences among treatments. Relative abundance ranges from blue (lower than the average percentage value) to red (higher than the average percentage value). The Compounds identified in the different apolar extracts derived from infested and non-infested roots were grouped in structurally related families (C).

Pot experiment
The pot experiments were conducted under controlled conditions to confirm field results and assess the underground infestation which was difficult to consider under field conditions. Data related to EON, NEO SDW and RDW recorded for different tested genotypes showed that the growth and biomass production have decreased in infested plants for all the genotypes compared to the non-infested plants.
These results showed that out of the five tested genotypes, only two genotypes ILL6415 and ILL7723 showed good resistance level under controlled conditions which confirm the results observed under open field conditions. Such resistance was expressed by the lowest shoot and root dry weight reduction and a low number of total O. crenata shoots and tubercules. similar results were reported in previous studies performed on faba bean and lentil (Ennami et al. 2017;Abdallah et al. 2020) who reported a large variation in infestation intensity under pots conditions and field trials.

Qualitative untargeted metabolomic analysis
In this part of the study, the two resistant genotypes ILL 6415 and ILL7723 with the susceptible genotype Zaaria were selected to conduct a qualitative metabolomic analysis to explore the biochemical mechanisms involved in the resistance against O. crenata. The principal components analysis (PCA) is commonly used in non-targeted metabolomics studies to perform metabolite correlation networks (Arbona et al. 2013). Based on PCA for shoot metabolites performed for both resistant genotypes ILL6415 and ILL7723 and compared to the control samples Orobanche parasitism was found to have a less effect on metabolites changes compared to the susceptible genotype Zaaria ( Figure  4(B)). However, the parasitism was found to have a markedly effect on metabolites change in the root metabolite for all genotypes compared to their respective noninfested control plants ( Figure 5(B)). The changes in metabolomic profile was reported also by Clermont et al. (2019) between the facultative parasite T. versicolor and the obligate holoparasite P. aegyptiaca with the host species M. Truncatula and A. thaliana, respectively. Similarly, other authors reported that the parasitism by P. aegyptiaca and O. foetida has affected respectively tomato (Amir 2016) and faba bean (Trabelsi et al. 2017;Abbes et al. 2020) metabolites when compared to non infested plants.
The results demonstrated that O. crenata parasitism may modify the shoot and the root metabolites of the host plants, especially for susceptible genotypes. Indeed, out of the detected metabolites, the increase of α-linolenic acid in the shoot of the resistant genotype could be related to a specific biochemical pathway of resistance, this essential fatty acid is an important component of the cell membrane that maintain the membrane integrity and functionality, prevents the membrane rigidification under stress conditions and enhance the production of ROS via the activation of Ca 2 + -ATPase (Delavault et al. 2017;Jia et al. 2020). Besides, αlinolenic acid is a precursor of jasmonic acid (JA) synthesis, a key rules in mediating resistant responses of plants under biotic and abiotic stresses by eliciting the production of alkaloid, terpenoid, coumarins, phytoalexins and taxane compound that are widely known by their functional value to plant under stress and regulating plant growth and development (Delavault et al. 2017). However, the accumulation of arachidic acid in the shoot and the root of the resistant genotype ILL6415 may serve as a defensive agent to fight the parasite attack, because several phytohormones and secondary metabolites are derived from this compound (Scharenberg et al. 2019). This long-chain fatty acid could be related to the wax, cutin and suberin production pathways, which help the plant to reduce water loss and protect the plant surface (Mutale-joan et al. 2020). Similar accumulation in α-linolenic acid and arachidic acid were reported by Jia et al. (2020) in drought-tolerant Populus simoniicv and drought-susceptible P. deltoides under drought conditions stress. Otherwise, clear difference was observed in root sterol content between the resistant and susceptible genotypes and between infested and non-infested plants. Wang et al. (2012) suggested that plant cells involve mechanical defense using sterols to make a physical barrier against the nutrient efflux to prevent nutrient and loss against bacterial and pathogens attack. In contrast to previous studies that focused more on polar metabolite variations including starch, amino acid and some secondary metabolite in response to parasitism in different plants (Abbes et al. 2009;Amir 2016;Clermont et al. 2019), the current study put the light-on fatty acids and sterols as important metabolites that can serve as regulatory pathways to conduct defense mechanisms in lentil and support the suggestion of the ability of the host plant to regulate phloem composition depending onthe host-parasite interaction (Jokinen and Irving 2019). Further, much remains unknown about the host-parasite biochemical interactions and a lot of work is still needed in exploring novel metabolic pathways and associated gene expression, genomic and proteomic analysis.

Conclusion
The broomrape O. crenata is one of the major problems limiting the production and the development of lentil in Morocco and many other countries in the region. Identification and development of resistant germplasm remains the best option to control this parasite. In this study, potential sources of resistance were identified out of 80 genotypes screened and evaluated under high O. crenata infested field and controlled conditions. Both genotypes ILL6415 and ILL7723 showed the highest level of resistance to the crenate broomrape. Such resistance was associated with complex physiological and biochemical mechanisms such as an increase of some specific metabolite biosynthesis that contribute in improving the host immunity and fight against the parasite. These mechanisms, if combined with other potential physical and/or chemical mechanisms, through classical breeding and/or genetic engineering approches, could help in improving the resistance to O. crenata in lentil.

Acknowlegements
This work was undertaken as part of research activities of crop improvement program at Mohammed VI Polytechnic University (UM6P) with contribution from ICARDA and Moroccan Foundation for Advanced Science, Innovation and Research (MAScIR).

Disclosure statement
No potential conflict of interest was reported by the author(s).

Notes on contributors
Youness EN-Nahli is a Ph.D. candidate at African Integrated Plant and Soil Research Group (AiPlaS), University Mohammed VI Polytechnic (UM6P) -Morocco, the International Center for Agricultural Research in the Dry Areas (ICARDA) and Mohammed V University -Rabat. He is conducting his research activities on screening and evaluation of lentil germplasm against broomrapes, identification of potential resources of resistance and associated genes/QTLs and investigation of physiological and biochemical mechanisms involved such resistance and genes-pathway's expression.
Hicham El Arroussi, Senior Scientist, R&D Manager of physiology and biotechnology of microalgae at Moroccan Fondation for Advanced Science, Innovation and Research (MAScIR). Affiliate professor at Mohammed VI Polytechnic University (UM6P). He has expertise on natural resources, microalgae production and valorization in different applications such as applied environment, agriculture and bioenergy. Shiv Kumar Holds a Ph.D. in Plant Breeding from the GB Pant University of Agriculture and Technology, Pantnagar, India. His area of expertise includes crop improvement through resistance breeding, widening the genetic base through pre-breeding, and development of genetic and genomic resources in food legumes. He is leading ICARDA's Food legumes program which aims to deliver improved germplasm of lentil, kabuli chickpea, faba bean and grass pea to national partners in South Asia, Sub-Saharan Africa, West Asia, and North Africa. He works on developing short duration climate smart varieties of lentil and grass pea with high iron and zinc content for sustainable intensification of cereal based cropping systems.
Outmane Bouhlal is a Ph.D. candidate at the International Center for Agricultural Research in the Dry Areas (ICARDA) and the University of El-Jadida -Morocco. He is working on nutritional quality and drought tolerance in barley.
Rachid Mentag holds a PhD in plant biotechnology from University Laval -Canada. He has expertise in plant biotechnology and molecular biology, Plant pathology and Plant-pathogen interaction. Actually, he is working as senior research scientist at INRA-Morocco with research focus on host genetic diversity of food legume crops including faba bean, lentil and chickpea. He has expertise on parasitic weeds management in legume crops with more focus on parasite-host interaction and host specificity.
Kamal Hejjaoui Ph.D student conducting research on lentil breeding at the international Center for Agricultural Research in Dry Areas (ICARDA) with focus on screening and identification of superior high yielding lentil germplasm with resistance to heat and drought.
Ahmed Douaik currently works at the National Institute of Agricultural Research (INRA-Morocco). He is skilled in Statistics, Spatial Analysis, and Soil Science. He holds a BSc in Agricultural Sciences from Hassan II Institute of Agricultural Sciences and Veterinary Medicine (IAV-Hassan II), Morocco, an MSc in Biometrics from University of Liège, Gembloux, Belgium, and a PhD in Space-Time Statistics from Ghent University, Ghent, Belgium.
Zouhaier Abbes holds a PhD in biology from Université de Tunis El Manar -Tunisia and University of Nantes -France. He is a plant physiologist skilled in parasite-host plant interaction and identification of potential biochemical, physical and physiological resistance mechanisms. he has also expertise in agricultural, genetics, biochemistry, molecular biology and environmental science. He is currently working at Higher Institute of Sciences and Technology of Environment, Borj Cedria and National Institute of Agricultural Research of Tunisia.
Nour Eddine Es-Safi holds a PhD in organic chemistry from Mohammed V University in Rabat, Morocco. He undertook postdoctoral at INRA-France. He is currently working at Mohammed V University, Rabat-Morocco. His research focuses on natural products and their structural elucidation, biological activities, and role in food technology.
Moez Amri holds a PhD in biological sciences and Plant breeding and HDR in Agricultural Science from the University of Carthage -Tunisia. He is an international scientist skilled in plant breeding with proven record in genetic enhancement of food legume crops as and their integration into the cropping systems for healthy soils and better nutritious food. His area of expertise includes crop improvement in particular food legumes breeding and development of superior high yielding germplasm with resistance to major biotic and abiotic stresses. He is also conducting research activities on parasitic weeds management in field crops with more focus on genetic resistance and parasite-host interaction.