Intercropping, which grows at least two crop species on the same pieces of land at the same time, can increase grain yields greatly. Legume-grass intercrops are known to overyield because of legume nitrogen fixation. However, many agricultural soils are deficient in phosphorus. Here we show that a new mechanism of overyielding, in which phosphorus mobilized by one crop species increases the growth of a second crop species grown in alternate rows, led to large yield increases on phosphorus-deficient soils. In 4 years of field experiments, maize (Zea mays L.) overyielded by 43% and faba bean (Vicia faba L.) overyielded by 26% when intercropped on a low-phosphorus but high-nitrogen soil. We found that overyielding of maize was attributable to belowground interactions between faba bean and maize in another field experiment. Intercropping with faba bean improved maize grain yield significantly and above-ground biomass marginally significantly, compared with maize grown with wheat, at lower rates of P fertilizer application (<75 kg of P2O5 per hectare), and not significantly at high rate of P application (>112.5 kg of P2O5 per hectare). By using permeable and impermeable root barriers, we found that maize overyielding resulted from its uptake of phosphorus mobilized by the acidification of the rhizosphere via faba bean root release of organic acids and protons. Faba bean overyielded because its growth season and rooting depth differed from maize. The large increase in yields from intercropping on lowphosphorus soils is likely to be especially important on heavily weathered soils.intercropping ͉ interspecific rhizosphere effect ͉ overyielding I ntercropping, which is the intermingled growth of two or more crops, with Ͼ28 million hectares of annually sown area in China (1), is also common in other parts of the world, such as in India, Southeast Asia, Latin America, and Africa (2). On nitrogen-deficient soils, legume-grass intercrops are known to overyield because of legume nitrogen fixation (2-5). About a third of terrestrial soils have insufficient available phosphorus (P) for optimum crop production, with many tropical acid soils being highly P-deficient (6, 7). Some pot experiments have suggested that legume/cereal mixtures can achieve greater P uptake on such soils (8-10) than can either species by itself. In field conditions, similar greater P uptake by intercropped maize with faba bean also was observed (11). However, both pot experiments and field experiments did not distinguish that the greater P uptake was derived from niche (rooting depth or seasonality) complementary or direct interspecific facilitation. We hypothesize that overyielding of intercropped species on P-deficient soil may result from a plant's chemical alteration of the rhizosphere that mobilizes P and thus enhances its own productivity and that of another species. We call this phenomenon the interspecific rhizosphere effect. Such chemical mobilization of a limiting nutrient would represent a mechanism of interspecific facilitation that, in concer...

SummaryDespite increasing evidence that plant diversity in experimental systems may enhance ecosystem productivity, the mechanisms causing this overyielding remain debated. Here, we review studies of overyielding observed in agricultural intercropping systems, and show that a potentially important mechanism underlying such facilitation is the ability of some crop species to chemically mobilize otherwise-unavailable forms of one or more limiting soil nutrients such as phosphorus (P) and micronutrients (iron (Fe), zinc (Zn) and manganese (Mn)). Phosphorus-mobilizing crop species improve P nutrition for themselves and neighboring non-P-mobilizing species by releasing acid phosphatases, protons and/or carboxylates into the rhizosphere which increases the concentration of soluble inorganic P in soil. Similarly, on calcareous soils with a very low availability of Fe and Zn, Fe-and Zn-mobilizing species, such as graminaceous monocotyledonous and cluster-rooted species, benefit themselves, and also reduce Fe or Zn deficiency in neighboring species, by releasing chelating substances. Based on this review, we hypothesize that mobilization-based facilitative interactions may be an unsuspected, but potentially important mechanism enhancing productivity in both natural ecosystems and biodiversity experiments. We discuss cases in which nutrient mobilization might be occurring in natural ecosystems, and suggest that the nutrient mobilization hypothesis merits formal testing in natural ecosystems.

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Even though ecologists and agronomists have considered the spatial root distribution of plants to be important for interspecific interactions in natural and agricultural ecosystems, few experimental studies have quantified patterns of root distribution dynamics and their impacts on interspecific interactions. A field experiment was conducted to investigate the relationship between root distribution and interspecific interactions between intercropped plants. Roots were sampled twice by auger and twice by the monolith method in wheat (Triticum aestivum L.)/maize (Zea mays L.) and faba bean (Vicia faba L.)/maize intercropping and in sole wheat, maize, and faba bean up to 100 cm depth in the soil profile. The results showed that the roots of intercropped wheat spread under maize plants, and had much greater root length density (RLD) at all soil depths than sole wheat. The roots of maize intercropped with wheat were limited laterally, but had a greater RLD than sole-cropped maize. The RLD of maize intercropped with faba bean at different soil depths was influenced by intercropping to a smaller extent compared to maize intercropped with wheat. Faba bean had a relatively shallow root distribution, and the roots of intercropped maize spread underneath them. The results support the hypotheses that the overyielding of species showing benefit in the asymmetric interspecific facilitation results from greater lateral deployment of roots and increased RLD, and that compatibility of the spatial root distribution of intercropped species contributes to symmetric interspecific facilitation in the faba bean/maize intercropping.

Plant diversity in experimental systems often enhances ecosystem productivity, but the mechanisms causing this overyielding are only partly understood. Intercropping faba beans (Vicia faba L.) and maize (Zea mays L.) result in overyielding and also, enhanced nodulation by faba beans. By using permeable and impermeable root barriers in a 2-y field experiment, we show that root-root interactions between faba bean and maize significantly increase both nodulation and symbiotic N 2 fixation in intercropped faba bean. Furthermore, root exudates from maize promote faba bean nodulation, whereas root exudates from wheat and barley do not. Thus, a decline of soil nitrate concentrations caused by intercropped cereals is not the sole mechanism for maize promoting faba bean nodulation. Intercropped maize also caused a twofold increase in exudation of flavonoids (signaling compounds for rhizobia) in the systems. Roots of faba bean treated with maize root exudates exhibited an immediate 11-fold increase in the expression of chalcone-flavanone isomerase (involved in flavonoid synthesis) gene together with a significantly increased expression of genes mediating nodulation and auxin response. After 35 d, faba beans treated with maize root exudate continued to show up-regulation of key nodulation genes, such as early nodulin 93 (ENOD93), and promoted nitrogen fixation. Our results reveal a mechanism for how intercropped maize promotes nitrogen fixation of faba bean, where maize root exudates promote flavonoid synthesis in faba bean, increase nodulation, and stimulate nitrogen fixation after enhanced gene expression. These results indicate facilitative root-root interactions and provide a mechanism for a positive relationship between species diversity and ecosystem productivity.flavanoids | gene expression | intercropping | root-root interactions | signals M any ecosystems, including grasslands (1, 2), forests (3), phytoplankton communities (4), and cropping systems (5), show a positive relationship between plant diversity and ecosystem productivity. Several mechanisms have been proposed to explain this relationship. A "sampling effect" occurs, because more diverse mixtures have a higher probability of containing a species with higher productivity (6). Complementarity effects occur when species vary in resource use and niche differentiation in space or time, leading to greater resource utilization (6-9). Facilitation occurs when one species increases the growth of other species through a wide range of processes (10). Facilitative effects may be direct (e.g., by shade or protection from harsh conditions) or indirect (e.g., when one species reduces attack by pathogens or herbivores on other species) (11-13).Legume/cereal intercropping systems have been widely studied in the context of diversity and ecosystem function and commonly overyield, because dinitrogen (N 2 ) fixation by legumes increases ecosystem nitrogen (N) supply (7,8), an example of facilitation. This facilitation is important to agriculture on a large scale, because app...