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CROP BREEDING, GENETICS & CYTOLOGY

Breeding Potential of European Flint and U.S. Corn Belt Dent Maize Populations for Forage Use

^a Centro de Investigaciones Agrarias de Mabegondo (CIAM), Apartado 10, 15080 La Coruña, Spain
^b IES Selgas, Avda. Selgas s/n, 33154 Cudillero, Spain

moreno_ciam{at}igatel.net

	ABSTRACT

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
Information about appropriate breeding populations and the type of forage maize (Zea mays L.) hybrids that should be developed for cooler regions of Europe is scarce. Our objectives were to determine the potential, performance, and heterosis of flint (F) and dent (D) populations as base germplasm for developing silage maize hybrids. Four U.S. Corn Belt dent populations which had been selected for early maturity and four European flint populations were included in our study. The eight populations, the 28 possible crosses among them, and topcrosses to inbreds A632 and EC18 were evaluated for stover, ear, and whole plant dry matter yield (DMY), as well as stover and whole plant dry matter content (DMC), digestible organic matter (DOM), and acid detergent fiber (ADF) content in northwest Spain. Means of F x D population crosses for stover and ear DMY (6.98 and 8.71 Mg ha^-1, respectively) were greater than those of D x D crosses (6.12 and 8.05 Mg ha^-1, respectively) and F x F crosses (6.03 and 7.64 Mg ha^-1, respectively). Average heterosis of whole plant DMY was greater for the F x D crosses (2.16 Mg ha^-1) than for the D x D and F x F crosses (0.23 and 0.56 Mg ha^-1, respectively). No significant heterosis was found for DOM and ADF of the stover and whole plant fractions. The only exception was a negative average heterosis of population BSSS-M for stover DOM. Results suggest that forage maize hybrids should be developed from the F x D heterotic pattern. Breeding strategies should include selection for stover DOM for both parent lines and hybrids because favorable recessive genes may be involved in this trait.

Abbreviations: ADF, acid detergent fiber • D, dent • DMC, dry matter content • DMY, dry matter yield • DOM, digestible organic matter • F, flint

	INTRODUCTION

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
IN COOLER REGIONS of central and western Europe, the use of maize as a forage crop has dramatically increased in the last three decades. Consequently, the metabolizable energy of the whole plant should now be considered for animal feeding purposes. To maximize selection advance for forage performance, the breeder has to consider both yield and quality traits of stover and grain simultaneously (Dhillon et al., 1990; Geiger et al., 1992). Hence, the major goals in forage maize breeding are improvement of whole plant dry matter yield, enhancement of feeding quality for ruminants, and suitable dry matter concentration of forage to ensure proper fermentation after ensiling (Struik and Deinum, 1990).

In contrast to the pure dent x dent crosses commonly employed in U.S. maize hybrids, most of the commercial hybrids grown in the cooler summer regions of central and western Europe are crosses between flint and dent inbreds. Several studies have shown that heterosis is exhibited in crosses between European flint and U.S. dent maize for grain and forage traits (Moreno-González, 1988a; Misevic, 1989, 1990; Ordás, 1991; Boppenmaier et al., 1992). These crosses combine local adaptation of the flint group with high yield potential of the dent lines. However, other traits such as lodging resistance (Moreno-González, 1988a) and dry matter content (Melchinger et al., 1992) do not fulfill desired requirements. Dhillon et al. (1990) reported that flint inbred lines transmitted more favorable digestibility related traits to the hybrid crosses than the dent inbred lines. Moreno-González et al. (1997) studied the performance and heterosis of flint and early dent germplasm for several traits and reported that dent germplasm exhibited less stalk and root lodging than flint germplasm. They concluded that because specific dent germplasm is being developed for short season areas, dent x dent grain commercial hybrids should progressively replace flint x dent hybrids in the cooler maize growing regions of Europe.

In spite of the genetic diversity represented by the U.S. and European maize accessions, the genetic base of the commercial hybrids both in the USA (Smith, 1988; Smith et al., 1992) and Europe (Smith et al., 1991; Messmer et al., 1992, 1993) is narrow. Among the potential consequences of this narrow genetic base are increased susceptibility to genetic shifts in pathogen populations and reduced rate of long-term genetic gain (Holland and Goodman, 1995). Therefore, it would be desirable to expand the genetic base of European cultivated maize by introduction of new germplasm and to guarantee a diverse supply of hybrids that minimizes the risk of genetic erosion (Smith, 1989). Evaluation of elite flint cultivars well adapted to western part of the Iberian Peninsula will also provide useful information for the improvement of flint x dent hybrids in this area.

The objectives of this study were to (i) evaluate European early flint populations and U.S. Corn Belt populations, which had undergone selection for earliness, for potential as base populations for developing forage maize hybrids, and (ii) determine the performance and heterosis of this germplasm for yield and forage quality traits.

	Materials and methods

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
Germplasm
Eight maize populations were studied: EPS6, SG1, `Aranga', and `Amarelo', having flint (F) endosperm, and SG2, BSSS-M, LANC-M, and BS10-M, having dent (D) endosperm. The origin and description of each population is detailed in Table 1 .

In the summer of 1990, the eight populations were topcrossed to the inbreds A632 and EC18 with pollen bulked from over 100 plants. The diallel crosses were carried out by pollinating about 50 plants from one population with 50 random plants from the other population. The crosses were made reciprocally, and seed from at least 80 pollinated plants from the two populations was bulked.

Experimental Design
The eight parental populations, their 28 diallel crosses, and the 16 topcrosses were grown in 1991 at three locations in northwestern Spain: Mabegondo (La Coruña), Puebla de Brollón (Lugo), and Pontevedra. The experimental design was an 8 by 8 triple lattice design that also included five adapted commercial hybrids as checks, the inbred testers A632 and EC18, the hybrid A632 x EC18, and other four genotypes not included in this study. Check hybrids were DEA, INRA 260, and Horreo 330, belonging to the maturity group FAO 200, and EVA, and Horreo 368, belonging to the maturity group FAO 300. The experimental unit was a one-row plot, 10 m long, and rows were spaced 0.8 m apart. Although they were grown at the same sites, these forage maize trials were independent from those already reported for grain maize at the same three sites (Moreno-González et al., 1997). Data collected were DMY (Mg ha^-1) of the whole plant, stover (including stalks, leaves, tassels and husks), and ear (including grain and cob) fractions; stover and whole plant DOM (g kg^-1); stover and whole plant ADF (g kg^-1); and stover and whole plant DMC (g kg^-1). Entries of each trial were harvested on the same date by machine at Mabegondo and by hand at the other sites. Representative samples of the stover fraction (approximately 1 kg), the whole plant (approximately 2 kg), and the ear fraction (3 ears) were taken from the harvested plants in each plot for determining the dry matter content and the nutritive value parameters. The stover and the whole plant samples were cut into 1-cm length pieces and oven-dried at 80°C for 18 h. The ears were also cut into 2-cm length pieces and oven-dried with forced air at 110°C for 18 h. The dried samples were later ground in a Christy and Norris 8'' mill (screen size = 1 mm). ADF and DOM were determined by the near infrared reflectance (NIR) equations from our CIAM laboratory for the stover and whole-plant fractions of maize. These equations were first calibrated with 100 samples by means of in vitro DOM (Tilley and Terry, 1963) and ADF (Goering and Van Soest, 1970) analysis, and later validated with 108 independent samples (Martínez, 1997).

The diallel crosses were further subdivided into those crosses involving different endosperm types [i.e., flint x dent (F x D) or dent x flint (D x F)], and those crosses involving the same endosperm type [i.e., flint x flint (F x F) or dent x dent (D x D)] (Table 2) .

where Y_kk' is the cross between populations k and k', µ_v is the mean of the eight parental populations, v_k and v_k' are the varietal effects of populations k and k', respectively, and h_kk' is the heterosis effect of the cross of populations k and k'. Furthermore,

where h is the overall average heterosis, h_k and h_k' are the varietal heterosis effects of populations k and k' respectively, and h_kk' is the specific heterosis effect of the population cross k x k'.

The average heterosis of a given population k across its seven (k') population crosses in the diallel (h_ak) was calculated as the mean of the seven h_kk' (k' = 1, 2, ...8, and k' k). The following expressions hold:

Note that s_kk' = 0, and h_k' = -h_k, for k' = 1, 2, ...n , and k' k, where n is the number of parental varieties (i.e., n = 8 ).

The average heterosis of groups F x F, F x D, and D x D were calculated as the differences between the means of the population crosses and the populations per se involved in the respective group (Moreno-González et al., 1997).

Errors associated with the overall average heterosis, the heterosis of each population averaged over all its population crosses in the diallel, the varietal heterosis effects and the average heterosis groups F x F, F x D, and D x D, were estimated according to Moreno-González et al. (1997). The error associated with s_kk' was estimated following Method 4 of Griffing (1956). Errors associated with the average heterosis of groups F x F, F x D, and D x D were calculated in similar way as the error associated with the overall average heterosis.

The analysis of variance of the 28 F₁ crosses and the eight varieties (populations) was performed following Analysis II of Gardner and Eberhart (1966). The adjusted means of the lattice designs in each environment were used in the analysis. Pearson correlation coefficients among some traits were obtained.

	Results and discussion

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
Dry Matter Yield
Populations Per Se
The flint population Aranga and the dent population BSSS-M had higher stover dry matter yield (DMY) than populations SG1, SG2, and Amarelo (Table 2). Population LANC-M had higher ear DMY than populations EPS6, SG1, SG2, and Amarelo. In addition, populations Aranga, BSSS-M, LANC-M, and BS10-M had better per se performances for whole plant DMY than populations SG1, Amarelo, and SG2. Therefore, there were significant differences among populations per se for the three DMY traits studied. The analysis of variance also revealed significant variety effects for ear, stover and whole plant DMY (Table 3) .

Diallel Crosses
All populations tended to have higher stover, ear, and whole plant DMY when crossed with populations having a different endosperm type than when crossed with populations having the same endosperm type (Table 2). The average DMY of population crosses in the F x D group was significantly higher than in both the D x D and F x F groups for the three yield traits (Table 4) . Koinuma et al. (1998) presented similar results, showing that hybrids with high dry-matter yields can be effectively developed by crossing dent with flint inbreds. This indicates that a productive heterotic pattern between F x D endosperm germplasm may exist for stover, ear, and the whole plant DMY. The same heterotic pattern was observed from the estimates of average heterosis between endosperm types (Table 4), which were significant and positive in the F x D group. Average heterosis estimates were also positive and significant in the F x F and D x D groups for ear DMY. These results were consistent with previous results for maize grain yield using the same germplasm (Moreno-González et al., 1997). However, heterosis between populations within groups (F x F and D x D) was not significant for stover and whole plant DMY. The percent average heterosis was always considerably higher for the F x D group than for the F x F and D x D groups for the three DMY traits (Table 4).

View this table: [in this window] [in a new window]   Table 6 Varietal and specific heterosis effects in the diallel crosses of eight maize populations for ear dry matter yield (DMY, Mg ha-1) above diagonal and for stover DMY (Mg ha-1) below diagonal

Topcrosses
There were no significant differences between the mean of topcrosses to flint inbred EC18 and that to dent inbred A632 (Table 2). However, flint populations tended to have lower stover, ear, and whole plant DMY when topcrossed to the flint inbred EC18, than did the dent populations. This is especially true for ear DMY in the topcross with Aranga, from which inbred EC18 was derived. This result again supports the heterotic pattern F x D. None of the evaluated populations topcrossed to EC18 had higher stover, whole plant and ear DMY than the A632 x EC18 hybrid. Thus, no population was superior to A632 when crossed with inbred EC18 for these three traits. Apart from the A632 x EC18 hybrid, the topcross of LANC-M population to A632 had the highest ear and whole plant DMY among all tested genotypes. The LANC-M x A632 topcross belongs to the Reid Yellow Dent x Lancaster heterotic pattern. Therefore, an effective breeding strategy would involve transferring favorable alleles from LANC-M population to EC18 to improve the A632 x EC18 hybrid.

When all genotypes (populations per se, population crosses, and topcrosses) were included in the analysis, ear DMY was correlated to stover DMY , but it was not correlated to the ear-to-stover ratio for DMY.

Digestible Organic Matter
On population per se basis, the flint population SG1 had the lowest digestible organic matter (DOM) for the stover, whereas the dent population BSSS-M had the highest (Table 2). The dent population BS10-M had the highest whole plant DOM among all populations and populations crosses; however, BS10-M had the lowest whole plant DOM among the topcrosses with inbred A632. On population per se basis, the range of variation was larger and the LSD was lower for stover DOM than for whole plant DOM. Thus, selection among populations should be more successful for stover DOM than for whole plant DOM.

There was no difference for stover DOM between the means of topcrosses to inbreds EC18 and to A632 (Table 2). However, some populations (e.g., LANC-M) tended to have a higher stover DOM when topcrossed to flint inbred EC18 than to A632, while some others (e.g., Amarelo, SG2, and BSSS-M) showed the opposite trend. These results suggest that the different performance of topcrosses for stover DOM does not depend on the different contribution of the two inbreds to the topcrosses, rather on the specific population-inbred combination.

Differences between topcrosses of single populations to inbred EC18 and A632 for whole plant DOM were not significant, except topcross BS10-M 3 EC18, which had higher whole plant DOM than BS10-M 3 A632.

Overall and group (F x F, F x D, and D x D) average heterosis was not significant for either stover or whole plant DOM (Tables 4 and 5). No population showed significant average heterosis for these traits, except BSSS-M, which had a negative average heterosis for stover DOM (Table 5). Furthermore, on a population per se basis, BSSS-M had also the highest stover DOM, which would indicate that DOM was not generally controlled by favorable dominant alleles in these populations. Therefore, effective breeding strategies should include selection for stover DOM for both the parents and hybrids to exploit possible additive and favorable recessive genes for this trait.

On population per se basis, there was a negative correlation coefficient between the ear-to-stover ratio and the stover DOM , whereas the correlation coefficient between the ear-to-stover ratio and the whole plant DOM was not significant . These results might be due to the dry matter translocation from the stover to the ear at harvest, which may be associated with reduction of digestibility in the stover fraction, but not in the whole plant. As the substantial effect of ear fill on stover composition should concern those evaluating stover quality of inbred lines (Coors et al., 1997), the ear-to-stover ratio should be also taken into account when selecting populations for stover quality.

ADF Content
There were no significant differences for ADF concentration between flint and dent population means on a per se basis; however, ADF differences among single populations per se were significant (Table 2). Dent population BSSS-M had less stover ADF than any other population, except Aranga. Flint population SG1 had the least ADF for the whole plant. There were not differences between means of topcrosses to inbreds A632 and EC18 for both stover and whole plant ADF; however, some specific topcrosses to EC18 (e.g., LANC-M 3 EC18) had less stover ADF than the counterpart LANC-M x A632. The opposite happened with population BSSS-M, where the topcross to EC18 had higher stover ADF content than that to A632. Thus, specific population-inbred combinations should be considered when selecting topcrosses for stover ADF.

ADF content was highly correlated to DOM for both the stover fraction and the whole plant , when all genotypes were considered in the correlation analysis. Either DOM or ADF can be used for determining quality parameters in forage maize. Correlation coefficients between ear-to-stover ratio and ADF were similar to those between ear-to-stover ratio and DMO, but with opposite sign.

Dry Matter Content
There were differences among both populations per se and population crosses for DMC of both the stover fraction and the whole plant (Table 7) . On population per se basis, the range of DMC for the whole plant (329 to 383 g kg^-1) was larger than for the stover fraction (224 to 257 g kg^-1) while the LSD values were similar for the two traits. The flint population SG1 had the highest stover and whole plant DMC on population per se basis, while dent population SG2 had the highest stover and whole plant DMC on population cross basis.

	Conclusions

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
The results concerning performance of populations, population crosses, and heterosis for ear DMY were consistent with previous results (Moreno-González et al., 1997) for maize grain yield. The additional results obtained for stover and whole plant DMY in this study should help the development of specific breeding strategies for forage maize hybrids. Among all the population crosses, those including flint x dent endosperm had the highest DMY on a stover, ear and whole basis. The ear fraction was mainly responsible for the heterosis expressed in whole plant DMY. The stover fraction contributed relatively little to whole plant DMY heterosis. Breeding strategies for silage hybrids should use populations from the F x D heterotic pattern. The Reid Yellow Dent x Lancaster heterotic pattern, after selection for earliness, also showed promise for the cool environments of Northwestern Spain for DMY. Populations differed for forage quality traits, but little heterosis was observed for these traits.

	ACKNOWLEDGMENTS

 
The authors thank two anonymous reviewers for valuable suggestions and assistance with English.

	NOTES

TOP NOTES ABSTRACT INTRODUCTION Materials and methods Results and discussion Conclusions REFERENCES

 
Research conducted at the CIAM and partially funded by the CICYT Grant AGF94-0301.

Received for publication August 5, 1999.

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