Heterosis and inheritance of drought tolerance in faba bean, Vicia faba L.

Twelve parental lines of faba beans and 27 F₁ hybrids derived from them were field-tested under a dry and a well-watered treatment, at two German locations (Göttingen, Hohenheim) in two years (1995, 1996). Drought was artificially induced using rain shelters from anthesis until maturity. Data were collected on yield under dry (Yd) and well-watered (Yw) conditions. Drought tolerance was assessed as Yd/Yw. The mid-parent heterosis was significant for Yd and Yw, but not for Yd/Yw. Relative heterosis for Yd (52.0%) was greater than for Yw (39.3%). Parental heritability was greater for Yw (0.86) than for Yd (0.61). However, it was similar for both traits in F₁ hybrids (0.65 and 0.66). Principal component analysis showed that F₁ hybrids exhibited the same pattern of adaptation as their parents. The heritability of drought tolerance was 0.48 in F₁ hybrids and 0.70 in parents. Attention should be paid to its negative correlation with yield. Several hybrids combined high tolerance, a favourable pattern of adaptation and appropriate performance.     

Hybrid performance and AFLP- based genetic similarity in faba bean

Successful prediction of heterosis and performance of F₁-hybrids from the genetic similarity of their parents based on molecular markers has been reported in several crops and can be very helpful in hybrid breeding. The relationship between genetic similarities based on amplified fragment length polymorphism (AFLP) of 18 European faba bean lines and their hybrid performance and heterosis was investigated. Parental lines, 62 F₁-hybrids and their F₂-progenies were evaluated in field trials in four environments in Germany for seed yield, 1,000-seed weight and plant height. Results clearly demonstrated a stable superiority of the hybrids over their inbred parents and elite check cultivars, and showed a marked and varying amount of heterosis. Parental seed yield and F₂-hybrid yield were promising as predictors for F₁-hybrids. AFLP analysis of the 18 inbred lines using 26 EcoRI/MseI primer combinations resulted in 1202 polymorphic fragments. Cluster analysis based on genetic similarity estimates unambiguously identified pedigree-related inbred lines. No clear separation of the 18 inbred lines into subgroups was detected. Correlation coefficients between genetic similarity estimates and either heterosis or F₁-hybrid performance were small and not useful. Also correlations between specific genetic similarity and specific combining ability were too small for all traits to be of predictive value. Results showed that AFLP-based genetic similarities are not useful to predict the performance of hybrids or heterosis within the elite European faba bean gene pool.     

Factors determining the performance of synthetics in Vicia faba L. 1. Heterogeneity,heterozygosity, and degree of cross-fertilization

Summary The relative performance of synthetic varieties in faba beans (Vicia faba L.) as compared with line varieties is the result of the joint effect of several factors. Factors investigated in the present study were (1) effects of heterogeneity and (2) heterozygosity on agronomic performance, (3) genetic variance for the degree of cross-fertilization among inbred lines and (4) genetic variance among homozygous and heterozygous genotypes for yield. The evaluation was based on multilocational trials in Germany with 36 inbred lines and derived entries with contrasting population structure (levels of heterozygosity and heterogeneity). When inbred lines and F₁-hybrids are included in the same mixture of genotypes, the resulting heterogeneity had a positive effect on yield. Relative mid-parent heterosis for yield was between 33% and 51%. The average degree of cross-fertilization was 53.5%, and its genetic standard deviation was 8.3%. The estimates for genetic standard deviation for yield among inbred lines were between 3.4 q/ha and 5.3 q/ha. The highest yielding synthetic variety is expected to outyield markedly the highest yielding line.     

Effectiveness of early generation selection under two plant densities in faba bean (Vicia faba L.)

The effectiveness of early generation selection under low and high plant density was evaluated in faba bean (Vicia faba L.). For this, the faba bean cv. 'Polycarpe' was pollinated by eight higher yielding cultivars of different origin. The F₁'s and F₂'s together with 'Polycarpe' and the pollinator with the highest yielding ability (cv. 'Effe') were evaluated under high plant density with adjacent controls and under low plant density in a honeycomb design. It was found that the eight hybrids had a different ranking at low than at high plant density for both generations (F₁and F₂). Hybrid Pol.x A-58 had the best combined yield performance in the F₁and F₂under high and hybrid Pol.x A-37 under low plant density. Application of individual plant selection in the F₂and F₃generation of the two crosses (Pol.x A-58 and Pol.x A-37), under both low plant density according to UNR-O honeycomb design and high plant density using a moving average, resulted in the selection of four groups with 20 selected F₃plants each. Within each group, equal numbers of seeds from each plant were mixed to establish four mixtures. These mixtures together with two controls ('Polycarpe' and 'Effe') were evaluated in a randomized complete block design (22 plants/m²) in one location for two years. It was found that the material selected continuously under low plant density had a significantly higher yield than the material selected under high plant density. In addition, the material selected from Pol.x A-37 (H₁R₂) under high plant density had a lower mean yield than that selected continuously under low plant density (H₁H₂). Their average yield over years was significantly higher than the yield of the material selected from Pol.x A-58 under both plant densities. Furthermore, the material selected from Pol.x A-58 under low plant density, had a significantly higher mean yield over years than its sibs selected under high plant density. It was concluded that honeycomb selection at low plant density can be used effectively for early generation selection in faba bean.     

Comparison of intra- and inter-pool crosses in faba beans (Vicia faba L.). I. Hybrid performance and heterosis in Mediterranean and German environments

Two recently detected, promising CMS systems in faba bean have opened up new possibilities in hybrid breeding. This study was conducted to determine hybrid performance and heterosis of 33 intra-pool and 66 inter-pool crosses from a factorial of six European small-seeded (Minor), six European large-seeded (Major), and eight Mediterranean faba bean lines. The parent lines and F₁ crosses were evaluated at two German and seven Mediterranean environments for anthesis, maturity, yield, yield components and plant height. Yield in the Mediterranean environments was 60% of that in the German environments. European Minor lines and their intra-pool crosses were unadapted to Mediterranean environments and yielded only 31 % compared with the Mediterranean lines and their crosses, whereas European Major genotypes yielded 58%. In German environments the European crosses yielded 107% of the Mediterranean crosses. Heterotic yield increase over the parental mean was largest for European Minor x Mediterranean inter-pool crosses, amounting to 95% in the Mediterranean and 73% in German environments. Our results confirmed the expectation of an increased heterosis in inter-pool over intra-pool hybrids and poor adaptation of European genotypes in Mediterranean environments. In the latter, Mediterranean intra-pool hybrids were the most promising, whereas in Germany inter-pool hybrids of the European Minor × Mediterranean type were most promising.     

Efficiency in the use of phosphorus,nitrogen and potassium in topless faba beans (Vicia faba L.) – variability and inheritance*

In a field experiment with topless faba beans, parental inbred lines and FI hybrids of a 7 × 7-diallel were evaluated for traits related to the P, N and K efficiency. In addition, the parental inbreds were cultivated in a pot experiment at two P levels (100 vs. 700 mg P per pot). Significant heterosis for grain yield, uptake and utilization efficiency of nutrients in the field experiment highlighted the enhancement of nutrient efficiency in the hybrids. Amongst both parental inbreds and hybrids, significant genotypic variation was found for nearly all recorded traits. In the diallel analysis, the GCA effects generally proved to be highly significant and in most cases considerably higher than the SCA effects. In the pot experiment, grain yield ranking of the parental lines at the high P level (= P700) was very similar to that in the field experiment, whereas at the low P level (= P100, – P deficiency) the ranking of the lines changed considerably. Correspondingly, the P level-line interaction was highly significant. High tolerance towards P deficiency was found for those two parental inbreds derived from cultivars bred under Syrian conditions. At the high P supply, P efficiency was more favoured by a high P uptake, but at P deficiency by a high internal P utilization.     

Genetic Structure of Three Open-Pollinated Faba Bean Varieties (Vicia faba L.)*

To study the genetic structure of open-pollinated faba bean varieties, three sets of genotypes were generated from each of the varieties ‘Minica’, ‘Kristall’, and ‘Deiniol’: (1) inbred lines developed by single-seed descent in bee-proof isolation cages, (2) intravanetal and (3) intervarietal F₁-hybrids produced by controlled hand crossing. In 1989 and 1990, a total of 144 entries, including the three open-pollinated source varieties, were grown in single-row plots with two replications at two locations in Western Germany. Performance data of the inbred lines revealed relatively large genotypic variability within each variety, for all the characters studied, which was greater in ‘Deiniol’ and ‘Kristall’ than in ‘Minica’. An average yield heterosis of 20 %, 39 % and 57 %, respectively, was found for the intravarietal hybrids, whereas that of the intervarietal hybrids varied between 70 and 73 %. Compared to the mean yield of the inbred lines the superiority of the open-pollinated source variety was small and not significant in ‘Minica’ (6 %), but it was large and highly significant in ‘Kristall’ (22 %) and ‘Deiniol’ (37 %). The present investigation revealed in faba bean that similar performance levels of open-pollinated varieties may result from either high per se performance under inbreeding with low heterosis values or from lower per se performance but with high heterosis values.     

Detection and characterization of two new CMS systems in faba bean (Vicia faba) *

Wide crosses were made to identify new cytoplasmic male sterility (CMS) systems in faba beans, based on the interaction of cytoplasm with restorer and maintainer alleles. A total of 330 F₁ hybrids were produced in both reciprocal forms. Male sterile segregates were observed in one reciprocal version in the F₂ generation of six crosses. Two of these crosses with female parents originating from Afghanistan and Egypt expressed stable male sterility in subsequent backcross generations. Based on the female parents of the two crosses, these two CMS systems were designated CMS 199 and CMS 297. CMS 199 was more stable than CMS 297 during backcross generations and across different environments. Maintainer and restorer lines for both CMS systems were identified. Lower expression of male sterility occurred in CMS 297 in the greenhouse during the winter generations than in isolation cages during the summer generations, which may be utilized to maintain male sterile lines by selfing. Regarding practical applications, the CMS 199 shows great promise for hybrid breeding in faba beans.     

Pollination, Fertilization and Seed Development in Inbred Lines and F1 Hybrids of Spring Faba Beans (Vicia faba L.)

When the flowers are not disturbed, the yield of F₁ hybrids of faba beans normally exceeds that in their inbred parents. The basis of this “heterotic autofertility” was investigated by examining flowers for pollination and fertilization throughout the flowering season and assessing the distribution of seeds and pods on mature plants. Six inbred lines and seven of their F₁ hybrids were grown without protection from bee visitation, and their autofertility was estimated’ by comparing their fertilization with that of cytoplasmic male-sterile line. The first direct evidence was obtained that heterotic autofertility acts by increasing; the proportion of flowers which were pollinated and fertilized. The autofertility was not perfect, especially in a closed-flower hybrid, as many ovules remained unfertilized. Several factors contributed to the number of seeds per pod in F₁ hybrids, being superior to parental means: (1) more ovules per flower, (2) greater fertilization of the basal ovules, (3) reduced abortion of the apical ovules and (4) greater overall maturation ovules.     

Heterosis for yield and other agronomic traits of winter triticale F1 and F2 hybrids

Triticale is generally treated as a self‐pollinating crop and line breeding is practised. Hybrid breeding has been discussed for some time, but there is little information for winter triticale. This study investigated heterosis for eight agronomic traits in F₁ and F₂ hybrids grown together with their parents as drilled plots in three environments. On average, grain yield heterosis was 12.5 dt/ha (a relative 10.5%) compared with the mid‐parent value for F₁ hybrids, and 6.2 dt/ha (5.0%) for F₂ hybrids and withawide range of 4.4–17.1 dt/ha for F₁ hybrids. A positive contribution to the heterosis of yield was made by kernels/spike and 1000‐kernel weight, whereas spikes/m² showed negative heterosis. Hybrid plants in F₁ and F₂ were taller than mid‐parents (8.3 cm and 5.3 cm, respectively), with a tendency to earlier heading. The negative heterosis for falling number in F₁ and F₂ hybrids could be a problem for commercial production of triticale hybrids. By selecting parents for combining ability and the identification of heterotic patterns, grain yield heterosis of 20% appears feasible.     

Dry Peas (Pisum sativum L.) Grown in Mixtures with Faba Beans (Vicia faba L.)—A Rewarding Cultivation Alternative

Two semi-leafless pea cultivars and two highly lodging-resistant faba bean cultivars were grown in a series of replacement trials conducted on a highly fertile loam soil near Gottingen from 1992 to 1995. Cultivation as well as harvesting of mixed crops of both species proved possible. In spite of the delayed harvest of the mixed crop stands determined by the later pod ripening of the faba beans (varying between seasons from -f 5 to +21 days), noticeable seed losses due to pod shattering were not observed for the overripe peas. Standing ability of the peas was considerably improved by mixed cropping. Lodging began later and was less severe at harvest. Within mixtures with 50 % or more faba bean proportions at sowing (relative to the seeding rate of the pure stands) the peas, n i most cases, remained nearly upright until harvest and instead of the usual 'Plattlager', i.e., prostrate but upright top, frequently a more highly inserted 'Hohllager', i.e., upright but collapsed at higher nodes, was observed. In contrast to these remarkably synergistic effects for standing ability, such effects proved to be only small and nonsignificant for grain yield (−1.5–2.9%). Yield stability of the mixed crops was higher than that of the pure stands of both species. Estimates of the stability parameters, ecovalence, deviation from regression and environmental variance were at least halved. Yield proportions of peas and faba beans in the harvested seed of the mixtures did not correspond with the seed proportions at sowing. They differed considerably between seasons and cultivar combination. Higher yield proportions of faba beans were observed under wet and cold weather conditions and for the cultivar combination Victor/Disco, whereas higher yield proportions of peas were found under dry and warm weather conditions and for the cultivar combination Mythos/Baroness .     

Indica-Japonica differentiation of paddy rice and its relationship with heterosis

The Indica‐Japonica differentiation of three photoperiod‐sensitive and/or thermosensitive genetic male‐sterile rice (PGMS or TGMS, respectively) lines and 47 male parental lines from seven ecotypes were studied for their restriction fragment length polymorphism marker data to determine which ecotype crosses with the three PGMS and/or TGMS lines could lead to higher yield potential, and to estimate the relationship between the Indica‐Japonica differentiation of parents and heterosis in grain yield and its components. The results indicated that hybrids derived from ‘N422s’ and the early‐middle ripening Indica varieties from southern China, and hybrids between ‘Pei'ai64s’ and three Japonica ecotypes, including North‐eastern Japonica varieties, restoring lines of Japonica hybrid rice and north China Japonica varieties, showed the highest grain yields. There was less variation of yield among the F1s between ‘108s’ and the seven ecotypes than among the other F₁s. Highly significant positive correlations between heterosis of the F₁ yield and genetic distance of the parents were detected, although the correlation between F₁ yield performance and genetic distance did not reach a significant level. Considerable variation of correlation between heterosis and genetic distance was also detected in the Indica × Indica crosses and Indica × Japonica crosses. There was much higher correlation (r = 0.63) between the F₁ yield performance and the genetic distance of parents in the Indica × Indica crosses than in the others. It is proposed that a genetic distance of 0.4‐0.8 between the two parents of hybrid rice might be appropriate not only for F₁ performance, but also for heterosis.     

Extent, variation and breeding impact of natural cross-fertilization in German winter faba beans using hilum colour as marker

Faba bean is a partially allogamous grain legume, mostly sown as spring crop in Central Europe, and in few mild coastal areas of UK and France as a winter crop. A population with a genetically wide base, conceived and selected for superior winter hardiness showing promise as winter crop in Germany was evaluated. The reproductive behaviour of this population was evaluated for its degree of cross-fertilization at five German environments (1999–2001), and inbred lines bred from it, plus one F1 bulk and further checks, were assessed for variation in cross-fertilization at three German environments (2001). Seed hilum colour, a dominant-recessive, biallelic locus, was used as a morphological marker. Cross-fertilization (population mean value of 60%) was shown to be higher than in spring faba beans and was environmentally influenced. Genetic variation was demonstrated, heritability was high (h²= 0.75), and heterosis was negative. Inbred lines with a cross-fertilization of 70% can be easily bred. Breeding synthetic cultivars is recommended, their optimum number of components is expected to be higher than the optimum recommended for spring faba beans. This revised version was published online in July 2006 with corrections to the Cover Date.     

A diallel analysis of various traits in winter triticale

In triticale, preharvest sprouting tolerance is important for yield stability and quality of the grain. Therefore, an experiment was conducted to estimate quantitative genetic parameters of preharvest sprouting tolerance in triticale using a diallel of eight modern triticale cultivars and breeding lines. The 28 F₁‐hybrids and the parents were planted in the breeding nurseries in 2001 and 2002 at four locations. Heading, plant height, falling numbers of two harvest dates, thousand kernel weight (TKW) and preharvest sprouting (Spr) were measured. The heritabilities for plant height and TKW were the highest with 0.91 and 0.85, respectively. Falling number and sprouting showed heritabilities between 0.66 and 0.83. Mid‐parent heterosis (MPHET) was negative for heading, indicating a non‐significant earlier heading of the F₁‐hybrids. In plant height and TKW the heterosis was positive with 4.7% and 6.3% MPHET. Both falling numbers displayed a negative MPHET of nearly −15%. In contrast to falling numbers, the preharvest sprouting of hybrids was slightly reduced in comparison with the parents. In all traits analysed, general combining ability (GCA)‐variances were greater than specific combining ability (SCA)‐variances.     

Genetic diversity in bread wheat, as revealed by coefficient of parentage and molecular markers, and its relationship to hybrid performance

Variable results have been obtained in different crop species using geneticsimilarity (GS) estimates based on molecular markers and coefficient ofparentage (COP) to predict heterosis.This study was designed: i) to assess the level of GS among 40 breadwheat (Triticum aestivum L.) cultivars selected in Central and SouthEurope; ii) to compare GS with COP; iii) to correlate the estimates ofparental diversity with the heterotic effects detected on 149 F₁hybrids obtained by crossing the 40 cultivars according to three matingdesigns.The parental cultivars and the F₁ hybrids were grown in severallocations at normal seed density. Significant heterotic effects were detectedfor grain yield and other traits including quality attributes. The parentalcultivars were assayed for DNA polymorphisms using two classes ofmarkers: 338 RFLP and 200 AFLP^® bands were scored. GS estimates werecomputed considering each molecular marker set separately (GS_RFLP,GS_AFLP) and together (GS_TOT). Ample differentiation amongthe parental cultivars was detected with the two marker sets. Although theaverage GS_TOT (0.43) was higher than COP (0.10), the twomeasurements were significantly correlated (r = 0.36, p < 0.01).Correlations between the different estimates of genetic diversity andF₁ performance or mid parent heterosis for grain yield and otherrelated traits were in general low although statistically significant.A more detailed analysis was conducted on 28 F₁ hybridsproduced in a half diallel cross of eight parental cultivars characterized byhigh heterotic effects for grain yield. The GS estimates based on RFLP,AFLP^® markers and also on RAPD were partitioned into general andspecific components. Correlations with general and specific combiningability effects for the measured traits were in several cases statisticallysignificant but too low to be predictive and therefore exploitable in practicalbreeding.     

Prospects for commercialisation of F1 hybrid field beans Vicia faba L.

Summary Yields of F₁ hybrids averaged 22% more than the mean of their heavier-yielding inbred parents, and the advantage of hybrids over composite populations varied from 3 to 18%, more in winter than spring beans.From estimates of cost/benefit ratios that included the possibility of poor seedset on male steriles, on average 1 year in 10, it was concluded that hybrid seed might cost more than twice that of conventional varieties, but a 10% increase in yield on a 4t/ha crop would cover this extra cost of seed.Thus, hybrids could still be the more profitable type of variety provided seed rates are low and yield levels high. It is therefore still worthwhile to pursue the goal of stable cytoplasmic male sterility (CMS) and/or a control of the rate of increase in revertants. Some methods are suggested whereby it might be possible to reduce revertants, improve pollination or exploit heterosis other than with CMS.     

Heterosis for yield and related characters in pea

Summary To determine the levels of heterosis in F1 hybrids, four current pea (Pisum sativum L.) cultivars from southern Australia were used as female parents and crossed with 18 introduced genotypes. The 22 parents, 72 F1 hybrids and, depending on the environment, either 54 or all 72 F2 families were grown in replicated plots in four environments. Grain yield, total dry matter, harvest index, branches per plant, pods per plant, seeds per pod, hundred seed weight, plant height, onset of flowering and flowering periods were evaluated. For both the F1 and F2 generation, heterosis was determined as the superiority over the mid-parent and also over the better parent. In addition, the superiority over the best commercial cultivar was calculated. Most hybrids were higher yielding than their mid-parent but were less stable in yield across environments. Four F1 hybrids were significantly higher yielding than the best parent, by up to 26%. There were significant correlations between F1 hybrid and mid-parent value for plant height, pods per plant and hundred seed weight but not for yield. Overall, grain yield heterosis was mainly due to more pods per plant in the hybrids. The level of heterosis for yield in a poor yielding environment was higher than that in a high yielding one. Both additive and non-additive gene effects were important in the expression of all studied traits. The average level of heterosis for grain yield and total dry matter in the F2 population was half of that in F1 hybrids. The low level of inbreeding depression from the F1 to the F2 generation suggested that epistatic gene action also contributed to the expression of grain yield. Some F2 populations maintained the high yield levels of the corresponding F1 hybrids.     

Heterosis performance of yield and fibre quality in F1 and F2 hybrids in upland cotton

Because of the difficulty of producing F₁ hybrid seeds by hand emasculation and pollination, wide use of heterosis in cotton production has been limited in China. The objective of this study was to evaluate the potential of F₂ hybrids for yield and fibre quality. A half diallel involving eight parents and their F₁ and F₂ hybrids was grown in replicated studies at Linqing and Nanjing in 1999 and Nanjing in 2000. Yield and fibre quality was determined for all 64 entries. Fibre quality was also determined for parents and F₁s, but only for Zhongmiansuo 28 (ZMS28), Xiangzamian 2 (XZM2) and Wanmian 13 (WM13) F₂s. These three F₂ hybrids are extensively planted in China and provide experimental controls with which to compare the performance of new hybrids. Average yield heterosis for F₁s and F₂s was 15.9 and 9.2%, respectively. Inbreeding depression for yield varied but some F₂s greatly out‐yielded the best variety. Average F₁ heterosis was 6.7, 6.2 and 2.9%, respectively for number of bolls per unit area, boll weight, and lint percentage. The average F₂ heterosis for the same traits was 4.4, 3.3 and 1.6%, respectively. F₁ heterosis for fibre traits was low. In general, parental average was a good indicator of the yield and fibre quality of F₁ hybrids. These encouraging results suggest there is sufficient heterosis for yield to use F₂s in China.     

Yield of F1, F2 and F3 hybrids of rice (Oryza sativa L.)

Summary Yield and yield components of F₁ hybrids were studied in three experiments at 30×30 cm spacings and in one experiment at 15×15 cm spacings. In the 30×30 cm experiments, 10 of the 41 hybrids tested significantly outyielded their high parents. However, only 2 hybrids significantly outyielded the best cultivar: one hybrid yielded 23 % and the other 16 % more than their respective check cultivars. The four hybrids in the 15×15 cm experiment yielded only 59 to 92% as much as their high parents.In areas where rice is transplanted at relatively wide spacings, the observed levels of F₁ heterosis in selected hybrids may be sufficient to warrant production of hybrid rice, if enough hybrid seed can be produced. For direct-seeding at the high rates normal in the USA, the relatively small levels of heterosis and the difficulties of hybrid seed production preclude use of F₁ hybrid rice cultivars at present.None of 19 bulk F₂ and F₃ hybrids in two experiments yielded significantly more than its high parent. Similarly, none of the 12 mixtures included in one experiment yielded significantly more than its high parent. On the basis of yield alone, using bulk F₂ or F₃ or simple mixture populations is not merited.Contribution from the Western Region, Agricultural Research Service, U.S. Department of Agriculture, and the Department of Agronomy and Range Science, University of California, Davis, California 95616.     

Effect of density,cultivar and irrigation on spring sown monocrops and intercrops of wheat (Triticum aestivum L.) and faba beans (Vicia faba L.)

Spring-sown intercrops of wheat (Triticum aestivum L.) and faba beans (Vicia faba L.) were grown in three experiments at the University of Reading, UK. One wheat cultivar, Axona, and one (experiment 1) or two bean cultivars (experiments 2 and 3) Scirocco and Maris Bead, were grown as sole crops and intercrops at 50%, 75% and 100% recommended density. Experiments were rainfed but irrigation was an additional treatment in experiment 3. Biomass and seed yields of both wheat and faba beans were greater when monocropped than when intercropped. There was no evidence that radiation use efficiency (RUE) of intercrops was significantly different from sole crops. In all intercrops there was no significant effect of density on biomass, RUE or seed yield, though there were compensating changes in yield components. Seed yields of Maris Bead were significantly greater than Scirocco in experiment 3 but not experiment 2. There was no significant effect of irrigation on RUE or on wheat biomass and seed yield, but there was a trend for irrigation to increase faba bean biomass (P = 0.07) and seed yield (P = 0.06). With later sowing in experiments 2 and 3, time to harvest was shorter and wheat and bean biomass, seed yield and RUE were reduced. All land equivalent ratio (LER) values for both biomass (maximum 1.23) and seed yield (maximum 1.44) were greater than 1, with one exception in experiment 3, indicating that intercrops of wheat and faba beans make more effective use of land than equivalent sole crops. Partial LERs for faba beans were always lower than those of wheat. The tendency was for highest LERs to occur at 75% recommended density.