World population is projected to reach over nine billion by the year 2050, and ensuring food security while mitigating environmental impacts represents a major agricultural challenge. Thus, higher productivity must be reached through sustainable production by taking into account climate change, resources rarefaction like phosphorus and water, and losses of fertile lands. Enhancing crop diversity is increasingly recognized as a crucial lever for sustainable agro-ecological development. Growing legumes, a major biological nitrogen source, is also a powerful option to reduce synthetic nitrogen fertilizers use and associated fossil energy consumption. Organic farming, which does not allow the use of chemical, is also regarded as one prototype to enhance the sustainability of modern agriculture while decreasing environmental impacts. Here, we review the potential advantages of eco-functional intensification in organic farming by intercropping cereal and grain legume species sown and harvested together. Our review is based on a literature analysis reinforced with integration of an original dataset of 58 field experiments conducted since 2001 in contrasted pedo-climatic European conditions in order to generalize the findings and draw up common guidelines. The major points are that intercropping lead to: (i) higher and more stable grain yield than the mean sole crops (0.33 versus 0.27 kg m −2 ), (ii) higher cereal protein concentration than in sole crop (11.1 versus 9.8 %), (iii) higher and more stable gross margin than the mean sole crops (702 versus 577€ha −1 ) and (iv) improved use of abiotic resources according to species complementarities for light interception and use of both soil mineral nitrogen and atmospheric N 2 . Intercropping is particularly suited for low-nitrogen availability systems but further mechanistic understanding is required to propose generic crop management procedures. Also, development of this practice must be achieved with the collaboration of value chain actors such as breeders to select cultivars suited to intercropping.

Grain protein concentration of durum wheat is often too low, particularly in low-N-input systems. The aim of our study was to test whether a durum wheat-winter pea intercrop can improve relative yield and durum wheat grain protein concentration in low-N-input systems. A 2-year field experiment was carried out in SW France with different fertilizer-N levels to compare wheat (Triticum turgidum L., cv. Nefer) and pea (winter pea, Pisum sativum L., cv. Lucy) grown as sole crops or intercrops in a row-substitutive design. Without N fertilization or when N was applied late (N available until pea flowering less than about 120 kg N ha −1 ), intercrops were up to 19% more efficient than sole crops for yield and up to 32% for accumulated N, but were less efficient with large fertilizer N applications. Wheat grain protein concentration was significantly higher in intercrops than in sole crops (14% on average) because more N was remobilized into wheat grain due to: i) fewer ears per square metre in intercrops and ii) a similar amount of available soil N as in sole crops due to the high pea N 2 fixation rate in intercrops (88% compared to 58% in sole crops).

we showed that intercropping of durum wheat and winter pea increased the yield and protein concentration of durum wheat when early N availability was less than 120 kg N ha −1 . The aim of the present work was to understand these results by analysing intercrop species dynamics for growth, light and N acquisition. A 2-year field experiment was carried out in southwest France with different fertilizer-N levels in order to compare wheat (Triticum turgidum L.) and pea (Pisum sativum L.) grown as sole crops and as an intercrop in a row substitutive design. The advantages of intercropping in low N conditions were due mainly to: (1) better light use (up to 10%), thanks to species dynamic complementarity for leaf area index and height; (2) growth complementarity over time (higher growth rate of wheat until pea flowering and then of pea until wheat flowering); and (3) dynamic complementary N acquisition associated with better wheat N status throughout growth. Disadvantages, underlining poorer complementarity within the intercrop stand, were observed with ample available N in early growth. This induced higher cereal growth during winter, which led to increase interspecies competition by reducing pea light absorption and, consequently, its biomass production.