Optimizing seminal sugarcane crosses: reciprocal effects and potential for superior familiesABSTRACT
Increases in sugarcane yield are achieved by the use of new Brazilian cultivars obtained by selection, cloning and/or crossing of superior materials. In this sense, the objectives of this study were to identify families with high potential for superior clone formation and to verify the reciprocal effect of seminal sugarcane parents. To this end, an experiment was conducted in an arrangement of Federer Blocks (2018/19 and 2019/20 harvests) with 40 families of full-sib, whose seedlings were obtained through seeds. The evaluated characteristics were Brix, plant height, stem diameter, and tons of sugarcane per hectare. The F07M43, F27M09, F29M28, and F40M39 families stood out together for the four traits studied. The existence of maternal and paternal effects in determining the performance of the progenies was evidenced. RB036088, RB036152, and RB966928 were identified as superior parents and represent promising options for improving yield. Sugarcane breeding programs should continue evaluating reciprocal effects to identify optimal parental combinations.
Key words
plant breeding; combining capacity; maternal effect; Saccharum spp.
INTRODUCTION
The expansion of sugarcane (Saccharum spp.) cultivation in recent years is mainly due to a well-established technological package and the adoption of new cultivars by producers, aiming to increase productivity and optimize the production of sugar, ethanol, and energy (Morais et al. 2015b, Berton et al. 2020, Carneiro et al. 2020, Oliveira et al. 2021).
In the genetic improvement of sugarcane, the processes of selection and cloning of superior genotypes of segregating populations predominate, which are obtained through crosses between different parents that present genetic variability (Morais et al. 2015b). In order to maximize this process, it is crucial to choose the appropriate parents and quantify the environmental effects of each trait to be selected (Morais et al. 2015a, Zhou 2021).
In the initial stages of sugarcane selection, when seedlings are used to obtain individuals, there is less efficiency in predicting breeding values (Bressiani 2002). This difficulty can be overcome by the use of mixed models, which allow simultaneous correction of environmental effects, in a large number of individuals, generations, and locations, generating more accurate estimates and predictions (Abu-Ellail et al. 2018, Hoarau et al. 2022).
Few studies have been conducted to elucidate the importance of parental contribution in determining progeny performance for major economically important traits in sugarcane. Zan et al. (2019) were among the first ones to investigate reciprocal effects in sugarcane crosses derived from seed rather than clonal propagation, demonstrating that reciprocal crosses can outperform their corresponding non-reciprocal combinations. Given the implications of the genetic effects arising from crossbreeding (Zhou 2021), particularly for the optimal choice of parents, studies addressing reciprocal effects are highly relevant to sugarcane breeding programs.
The development of methods that aid in the choice of parents can increase the chance of producing superior genotypes in given progeny (Hoarau et al. 2022). Diallel crosses are considered great options in this sense, due to the large amount of genetic information they can offer, such as inferences about hybrids and the combinatorial capacity of their parents (Cruz 2006, Fan et al. 2014, Zhou 2021), as well as important information about the reciprocal effects on different traits (Diniz et al. 2018, Wu et al. 1980, Zhou 2015).
It is noticed, however, that most of the studies deal with the study of family selection in sugarcane clones, and there are divergences regarding the inheritance of the traits (Lucius et al. 2014, Xavier et al. 2014, Santana et al. 2017). In addition, there is little literature available on parental studies and maternal effect on sugarcane (Shanthi et al. 2005, Zhou 2015).
There is, therefore, a need for new studies aimed at the selection of new parental genotypes, which in addition to being productive, bring together agronomic and quality characteristics (brix, sucrose, reducing sugars, fibers, minerals). In this sense, the present study aimed to identify families with high potential for superior clone formation and to verify the reciprocal effect of seminal sugarcane parents.
MATERIAL AND METHODS
The study was carried out, in the beginning, at the Serra do Ouro experimental station of the Inter-university Network for the Development of the Sugarcane Industry (RIDESA) and Universidade Federal de Alagoas (UFAL), Murici (9°18’26’’S, 35°55’55’’W and 450 m a.s.l.), Alagoas, Brazil, 2017. To conduct the field experiment, experimental biparental crosses were made using a total of 29 parents and resulting in 40 families of siblings, as shown in Table 1.
Sugarcane crosses and their reciprocal crosses and codes used to identify crosses used in the study of families composed of siblings in sugarcane series RB17. Maringá, Paraná state, Brazil, 2022.
As they are crosses carried out in 2017, these families belong to the so-called RB17 Series, of the Sugarcane Breeding Program, Universidade Federal do Paraná (PMGCA/UFPR)/RIDESA.
The seedlings were obtained from the seeds of the previously mentioned crosses on Table 1 and produced in the greenhouse of the Universidade Estadual de Maringá (UEM) Experimental Farm, in Iguatemi, Paraná state, Brazil, in 2018 and transplanted to the field in the same agricultural year. Seeds from each crosse were sown in trays containing commercial substrate and filter cake, using 3 g of seeds from each crossing package. Before transplanting to the field, a pre-selection was made in all trays to plant only the best developed seedlings. A total of 1,236 superior seedlings were selected, ranging from 10–40 individuals per family. Cultivars RB986928 and RB867515 were included as checks.
The field experiment was conducted in the 2018/2019 and 2019/2020 growing seasons at the Irrigation Training Center (CTI) of the UEM, located at 23º23’51” S, 51º56’56” W and alt 380 m a.s.l. The soil of the experimental area was classified as a dystroferric Red Latosoil (Embrapa 2018).
A Federer Blocks design containing 40 families (Table 1) with random repeats was adopted in the study. The treatments were allocated in 5-m long furrows spaced by 1.50 m between rows and 0.50 m between individuals. A minimum of 10 individuals were used per furrow. The cultural treatments and fertilization were carried out according to the technical recommendations for the sugarcane crop (Santos and Borém 2016).
The first harvest was carried out 14 months after planting, in 2019, and of ratoon cane, approximately 12 months after the first cut, in 2020. All harvests were carried out manually with the use of a machete, piling up the straw on the carriers.
The following agronomic characteristics were evaluated:
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Brix: expressed in degrees, obtained with the aid of a field refractometer with direct readings in °Brix of the stem juice. Three samplings were originated for each seedling, and the stalks were randomly sampled. The first measurement was taken in the lower third of the stalk, the second in the middle third, and the third in the upper third. At the end, the mean value of the three samples in each individual was considered;
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Plant height: expressed in meters, measured with a graduated ruler every 5 cm, totaling 4 m, which was placed in the center of the clump, and the height was measured up to the first visible auricle (dewlap), thus characterizing the +1 leaf;
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Stem diameter: expressed in millimeters with the use of a caliper. Three random stalks were chosen, and samplings were carried out in their middle third. The mean value of the three samples was considered;
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Biomass production in tons of sugarcane per hectare (TCH): calculated as a function of the average stem mass (M1C), the number of tillers per meter (NPM) and a correction factor as a function of the spacing (1.5 m), as described in the Eq. 1 (RIDESA 2009):
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The data obtained were submitted to restricted maximum likelihood analysis (REML) and best unbiased linear predictor (BLUP) using Eq. 2 of the Selegen REML/BLUP software (Resende 2007).
where: y: the data vector; r: the repetition effects vector (assumed to be fixed) added to the general average; a: the vector of individual additive genetic effects (assumed to be random); p: the vector of (random) plot effects; f: the vector of plot effects (random) full-sibling family dominance; b: the vector of (random) block effects, and “ε” : the vector of (random) errors or residuals.
Capital letters represent incidence matrices for the aforementioned purposes.
In obtaining the classic selection index (Smith 1936, Hazel 1943), all variables had the same weight. The Smith-Hazel has the very important advantage of being able to take account of inheritance and genetic and phenotypic correlations among traits in a way which should maximize the gain in total genetic merit (Cotterill 1985). The index is obtained through a linear combination of traits weighted by coefficients that are determined by considering the existing genetic and phenotypic covariances among the characters involved in simultaneous selection (Ramalho et al. 2024).
The parents were ordered by the classic selection index, standardized by the additive values of each parent, and the necessary procedures were also carried out by the Selegen—REML/BLUP program (Resende 2007).
RESULTS AND DISCUSSION
The rank of the superior families for the four studied characteristics is shown in Table 2. The results provide evidence that reciprocal effects can influence cross choice and family selection, in agreement with findings reported in other reciprocal effects studies, such as Zan et al. (2019). Among the advantages of the joint analysis, there are the greater accuracy and safety of the results in relation to genetic and environmental effects (Balsalobre et al. 2016). Between the 40 total families in the experiment, 23 elite families were selected for height (Table 2), 24 families for stem diameter (DIA), 19 families for BRIX and 21 families for TCH.
Classification of the best families based on genotypic values (Vg) from two harvests (plant cane and ratoon) of the 40 families of sugarcane siblings, Series RB17 for plant height (ALT) and stem diameter (DIA), BRIX and tons of sugarcane per hectare (TCH). Maringá, PR, Brazil, 2022*.
Taking into account the average of the selected families (µsel), it is expected that, with the selection of these families and the installation of the second phase of the UEM Sugarcane Breeding Studies, known as T2, the experimental average would increase by 4.74% (from 2.6217 to 2.7460) for ALT, 3.60% (from 25.4592 to 26.4115) for DIA, 4% (from 22.6563 to 23.6014) for BRIX and 11% (from 38.5283 to 43.3084) for TCH (Table 2).
At plant height, the first 12 families with high genotypic value were respectively F15M09, F01M14, F43M15, F09M15, F15M43, F02M31, F02M35, F28M29, F39M40, F40M39, F45M09, and F14M01. If this value is used to advance with these 12 families in the next selection phases of the UEM Sugarcane Breeding Studies, we can say that we would be increasing, on average, this trait by 13.8400% (7.6836 cm). A superior family is considered to be one whose relative average (%) generated by its selection adds values above 0 (Zeni Neto et al. 2019). As the reverse is true, families with relative means of Vgc ≤ 0 were disregarded, so Table 2 lists only the higher families.
Through the results presented in Table 2, it can also be observed that, when the next selection phase was installed with the 12 best families (µ12), considering Vg > 0 and µ above the overall mean, the experimental mean of ALT, DIA, BRIX, and TCH could be increased, respectively, to 2.7955, 26.8553, 23.907 ,and 44.5162, values that are relevant when compared only with the general average. Some studies have shown that the selection of better individuals becomes easier when families have higher genotypic values when compared to families with lower values (Lucius et al. 2014, Zeni Neto et al. 2019).
A low correlation was observed between the elite families during the studied seasons (ρ19/20) for ALT, DIA, BRIX, and TCH (0.5447; 0.0249; 0.1183; and 0.2641, respectively) (Table 2). Such values, even though they are of low correlation magnitude, are positive and contribute to the selection process. This allows us to infer that, on average in the ranking of the best families, a superior family in the 2018/19 harvest was also superior in the ranking of the 2019/20 harvest, even if in different placements. In other words, there is a trend of similar behavior in both crops studied. In this way, the need for joint analysis between the harvests is evidenced, so that the selection becomes more accurate.
The best family for ALT was F15M09, with a mean value of 2.8716 m, whereas, for stem diameter (DIA), F27M09 ranked first, with a mean of 29.1666 mm. For °Brix, F42M41 showed the highest mean (25.2878). Regarding TCH, reciprocal effect (maternal and paternal) was evidenced, as F35M02 exhibited the highest production, reaching 47.0128 t.ha-1, while its reciprocal cross, F02M35, did not appear among the best families (Table 2).
Another extremely relevant issue in relation to selection concerns the ranking of families according to each studied characteristic. It is difficult to find the same elite families for all the characteristics of importance for the improvement of sugarcane. This fact was also observed in the present study, in which for BRIX, for example, the best family was F42M41, which did not even appear among those selected for the other traits (ALT, DIA and TCH). Thus, if selected, the experimental average of these three characteristics would decrease. These results were also found by other authors in the sugarcane study (Xavier et al. 2014, Zeni Neto et al. 2019).
In view of the difficulty of selecting elite families for all traits, the strategy used by breeders is to try to select families that are average, that is, that even though they are not the elites, they are among those selected for all the traits evaluated in the breeding program. This was observed in the present study with the F07M43 family, which, even though it was not among the 12 elites, would be selected because it was among those selected for the four characteristics of the study at the same time (Table 2).
The F40M39 family deserves great attention, as it was among the 12 selected for three (ALT, DIA, and TCH) of the four traits simultaneously evaluated. In addition, the F27M09 and F29M28 families also showed superior performance, standing out among the elite for two traits evaluated and remained among those selected for the other traits (Table 2).
Table 3 shows the best parents that made up the RB17 series for the four traits in descending order of their additive values (α). Parents were selected for ALT, DIA, BRIX, and TCH. It is expected that these parents will contribute to the improvement of the traits under study in the next crosses.
Classification, based on additive values (α), and genetic gain (g%), from two harvests (plant cane and ratoon) for all parents used to originate the RB17 series through biparental crosses, RB17 Series, for plant height (ALT), stem diameter (DIA), BRIX and tons of sugarcane per hectare (TCH). Maringá, PR, Brazil, 2022.
As the joint analysis showed additive effects of the parents with the harvests for ALT, DIA, and TCH (8.8981, p < 0.01; 5.2476, p < 0.05 and 5.6888, p < 0.05 respectively), the ordering of the best parents varied according to the harvests. Such parents can be selected to compose the next stages of crossbreeding of the breeding program, without having so much influence from this differentiated ordering.
For the traits ALT, DIA, and BRIX, significant gains are expected in the next series of crosses (7.7905% RB036152 parent, 4.120% RB931530 parent, and 10.0569% SP80-3280 parent, respectively). Such gains may be linked to the greater genetic variability found in the population (Table 3). On the other hand, for TCH, this may indicate that the parental genomes for this trait share a similar genetic basis, which may limit the magnitude of genetic gains.
In the same way that occurs for the selection of the best families, in the selection of the best parents, it is also unlikely that an elite parent for a given trait will be an elite parent for all the other traits evaluated in the breeding program. This result is shown in Table 3, in which the two best parents for TCH (RB966928 and RB97319) did not appear in the lists of the best parents for ALT, DIA, and BRIX, because their genotypic average would not be sufficient to promote an increase in these three characteristics for future crosses of the UEM Sugarcane Breeding Studies.
Thus, the procedure adopted by the breeders in the selection of parents is the same as that adopted for the selection of families, that is, parents who are among the list of selected parents are preferred, even if they are not the first placed. In this context, the parents RB931530, RB946015, and RB936109 even though they are not among the 12 best for the four traits, should be chosen to compose RIDESA’s future new series. Besides, the parents RB996519 and RB996961 (Table 3) also stood out since they are among the elites for all traits evaluated simultaneously. Therefore, significant gains are expected in the traits evaluated in sugarcane.
It is important to emphasize that the results mentioned above are important to continue the improvement of sugarcane via cloning. However, such results are also extremely relevant in the selection of parents regarding gender in the composition of crosses.
A reciprocal effect was found in the materials evaluated, with variations in the characteristics depending on the parents, sometimes used as a female or as a male. Analyzing, for example, the RB946015 and RB936109 parents, considered promising in the selection process (Table 3), they were the ones that made up the F29M28 and F28M29 families (Table 1). Between them, F29M28 (Table 2) was among those selected for all the characteristics of this study. This indicates that the RB946015 parent should be used as a female and the RB936109 as a male, in the expectation of obtaining improvement in the characteristics (ALT, DIA, BRIX, and TCH) in future crosses.
Similarly, when using the RB996519 and RB996961 parents, both promising, the F40M39 family, the result of the crossing of both, stood out for the four characteristics studied. Thus, there is a reciprocal effect with advantages in additivity when the RB996961 parent was used as a female.
Another parent that showed reciprocal effect was RB931530 (Tables 1 and 3), because when crossed with RB036088, it gave rise to the F27M09 family (Table 2), which is among those selected to compose the new stage T2 of the UEM Sugarcane Breeding Studies.
Table 4 shows the best sugarcane parents ordered by the classic selection index standardized by the additive values of each parent. Selection indices help in choosing parents for future crossings (Hoarau et al. 2022). In this study, the same weight was considered for the four traits. Almeida et al. (2014) showed that the classical selection index allowed superior simultaneous gains in sugarcane traits, in comparison with others index. Besides, this index is widely used in other crops (Heinz et al. 2012, Jahufer and Casler 2015).
Ranking based on the classic index, from two harvests (plant and ratoon cane), for the four traits studied (ALT, DIA, BRIX and TCH) for the sugarcane parents that composed the RB17 Series. Maringá, PR, Brazil, 2022*.
When selecting and crossing, Table 4 shows that, for example, for the RB931530 and RB036088 parents, gains in the order of 0.291 to 0.287% could be expected for ALT, DIA, BRIX and TCH, simultaneously. The RB931530 parent, with the highest selection index (Table 4), showed favorable results for DIA and TCH, when used as a female in crosses with RB036088. Furthermore, superior performance for TCH was observed when RB036088 was used as the male parent in crosses with RB036152, as well as for ALT and DIA in crosses with TUC74-15.
For the parent RB996961 highlighted for the selection index, it made it possible to select families for ALT and TCH (Table 3), with advantages when used as a female in the cross with RB996519. While the RB975932 parent should be used as a female, as it also stands out in the selection of the best families for TCH.
The selection index once again demonstrated to be an important auxiliary tool for breeders, as it allows grouping results of the studied characters. This was explicit, for example, for the RB996519 parent, who was in fourth position (Table 4), but occupied the sixth (ALT), fourth (DIA), eleventh (BRIX) and fifth (TCH) positions in the individual classification of each variable (Table 3).
General combining ability (GCA) and the specific combining ability (SCA) were also analyzed in this study, as presented in Table 5. GCA reflects the average performance of genotypes across multiple hybrids combinations and is primarily related to additive genetics effects (Hallauer et al. 2010), while SCA relates to the non-additive portion of the total variance was predominantly influenced by dominance deviations and epistasis (Rojas and Sprague 1952).
Results of the deviance analysis for plant height (ALT), stalk diameter (DIA), BRIX and tons of sugarcane per hectare (TCH) obtained from two harvest (plant cane and ratoon cane) for 40 full-sib sugarcane families, RB17 Series. Maringá, PR, Brazil, 2022.
The joint deviance analysis demonstrated the presence of significant additive genetic effects (GCA) for the traits ALT, DIA and TCH (Table 5). In the case of ALT, significant GCA effects (χ2 = 8.9095, p < 0.01) were detected along with a significant GCA × harvest interaction (χ2 = 8.8981, p < 0.01), highlighting the relevance of additive genetic control and its variable expression across different harvests. Moreover, the signific of SCA effects for this trait indicates that both additive and non-additive genetic components contribute to its phenotypic expression.
For DIA, significant GCA effects were observed (χ2 = 5.2572, p < 0.05), in addition to the significant interactions involving SCA × harvest and GCA × harvest. These results suggest that the expression of this trait is governed by a combination of additive and non-additive effects, whose is modulated by environmental conditions.
In contrast, no significant effects were identified for BRIX, regardless for GCA, SCA or their interactions with harvest, suggesting limited genetic variability for this trait among the parents and crosses. With respect to TCH, significant GCA effects were detected (χ2 = 9.798, p < 0.01). Furthermore, a significant GCA × harvest interaction was observed, with χ2 = 5.6888 (p < 0.05), demonstrating the predominance of additive genetic effects and their sensitivity to environmental conditions. The lack of significant SCA effects for TCH supports the greater importance of additive control for this trait in the evaluated genotypes.
Estimates of mean family heritability (hˆ2fam) revealed contrasting genetic control among the evaluated traits. ALT and BRIX showed low heritability values, indicating a predominance of environmental influences and genotype × harvest interaction. Conversely, DIA and TCH exhibited moderate heritability, emphasizing the relevance of additive effects and suggesting favorable conditions for selection based on breeding values.
Considering the significant GCA effects detected, the discussion must focus on TCH as a representative trait, since it is the most important agronomic trait for sugarcane yield and is used to determine the price and payment method of sugarcane in Brazil (CONSECANA 2006). The estimation of general combining ability becomes essential to identify superior parents and guide crossing strategies in sugarcane breeding programs.
The estimates of general combining ability for TCH are presented in Table 6 and reveled substantial variation among the evaluated parents. Several parents exhibit significant and positive GCA values, indicating a favorable contribution of additive genetic effect to total sugarcane yield.
Estimates of general combining capacity (GCA) for tons of sugarcane per hectare (TCH). Maringá, PR, Brazil, 2022.
Among the evaluated parents, RB036088, RB036152, and RB966928 showed the highest positive estimates for TCH, highlighting their potential for use in crosses aimed at yield improvement. These results are consistent with previous studies in sugarcane, which have reported a predominance of additive effects for yield related traits (Bressiani et al. 2002, Barbosa et al. 2005, Xavier et al. 2014).
Although some parents presented lower GCA estimates for TCH, they should not be entirely discarded, as they may still contribute with valuable alleles when used in the formation of families or in specific breeding strategies, allowing the evaluation of their progenies in subsequent selection phases.
Differences between the present results and those reported by Yadav et al. (2021) and Zhou (2021), who observed a greater contribution of non-additive effects for TCH, may be attributed to differences in the genetic background of the parents, allele frequencies, particularly if self-fertilization occurred at any stage, genetic divergences and environmental conditions under which the studies were conducted.
The best parents were RB036088, RB036152, and RB966928, which should be further explored by breeding programs to increase the TCH characteristic. Another possibility would be to use these individuals as positive GCAs in the formation of a population so that the progenies can be evaluated in future phases of breeding programs.
CONCLUSION
The results highlight the importance of studying reciprocal effects in sugarcane, as differences were observed among crosses depending on whether a parent was used as male or female. These findings reinforce that parental role can influence the expression of key agronomic and industrial traits. Therefore, it is of great importance that sugarcane breeding programs continue to investigate reciprocal effects, aiming to identify the most suitable parental combinations. Understanding which parents perform best as male or female will allow breeders to make more informed crossing decision and to consistently use superior parents in future breeding cycles, thereby increasing the efficiency of selection for economically important traits in sugarcane.
ACKNOWLEDGMENTS
The authors would like to thank the Universidade Estadual de Maringá (UEM) and the Agronomy Graduate Program.
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How to cite:
Santos, R. F., Zeni Neto, H., Moterle, L. M., Borsuk, L. G. M., Montini, L. M., Cristiano, V. B., and Scapim, C. A. (2026). Optimizing seminal sugarcane crosses: reciprocal effects and potential for superior families. Bragantia, 85, e20250244. https://doi.org/10.1590/1678-4499.20250244
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FUNDING
Not applicable.
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DECLARATION OF USE OF ARTIFICIAL INTELLIGENCE TOOLS
The authors declare no use of artificial intelligence tool in the preparation of the manuscript.
DATA AVAILABILITY STATEMENT
All data were generated or analyzed in this study.
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Section Editor:
Gabriel Constantino Blain https://orcid.org/0000-0001-8832-7734
