Journal of
Plant Breeding and Crop Science

  • Abbreviation: J. Plant Breed. Crop Sci.
  • Language: English
  • ISSN: 2006-9758
  • DOI: 10.5897/JPBCS
  • Start Year: 2009
  • Published Articles: 410

Full Length Research Paper

Agro-morphological characterization of Fonio millet accessions (Digitaria exilis Stapf.) collected from Boukoumbé, Northwest of Benin

Emmanuel Sekloka*
  • Emmanuel Sekloka*
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Cyrille Kanlindogbe
  • Cyrille Kanlindogbe
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Samadori Sorotori Honore Biaou
  • Samadori Sorotori Honore Biaou
  • Laboratory of Ecology, Botany and Plant Biology (LEB), Department of Natural Ressources Management, Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Hubert Adoukonou-Sagbadja
  • Hubert Adoukonou-Sagbadja
  • Laboratory of the Genetic Resources and Breeding of Species, Department of Genetics and Biotechnology, Faculty of Sciences and Technologies, University of Abomey-Calavi, Benin.
  • Google Scholar
Albert Kora
  • Albert Kora
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Fidele Tchossi Motouama
  • Fidele Tchossi Motouama
  • Laboratory of Ecology, Botany and Plant Biology (LEB), Department of Natural Ressources Management, Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Moudjaidou Seidou
  • Moudjaidou Seidou
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Valerien Amegnikin Zinsou
  • Valerien Amegnikin Zinsou
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Leonard Afouda
  • Leonard Afouda
  • Laboratory of Phytotechny, Plant Breeding and Plant Protection (LaPAPP), Department of Sciences and Techniques of Vegetal Production (STPV), Faculty of Agronomy, University of Parakou, Benin.
  • Google Scholar
Lamine Baba-Moussa
  • Lamine Baba-Moussa
  • Laboratory of Biology and Molecular Typing in Microbiology, Department of Biochemistry and Cell Biology, Faculty of Sciences and Technologies, University of Abomey-Calavi, Benin.
  • Google Scholar


  •  Received: 20 June 2016
  •  Accepted: 05 September 2016
  •  Published: 31 October 2016

 ABSTRACT

Fonio is a cereal food of great socio-economic and cultural importance in south Sahara African Countries. Unfortunately, it is practically absent from National Agricultural Research Programs. To characterize the agro-morphological diversity of fonio ecotypes grown in Benin, twenty accessions collected from Boukoumbé were evaluated in a randomized complete block design (RCBD) with three replicates in Parakou. Significant variability was detected for several characters. The early accessions matured at approximately 90 days after sowing with yields below 800 kg/ha. The late accessions matured in 100 days and the most productive recorded more than 1.5 t/ha. Factor analysis of mixed data helped to classify the accessions into four morphological groups. Chi-square independence test showed that collar color, green color of foliar limb, anthocyanin coloration and its distribution in different aerial organs, type of panicle and panicle exsertion were the most discriminating qualitative parameters. The λ-wilk test revealed that date of flowering, plant height, length of panicle leaf, length of racemes and grain yield were the most discriminating quantitative traits. This study enabled a better knowledge of cultivated ecotypes and distinguishing criteria. The variability observed offered interesting perspectives for genetic progress through breeding programs of these ecotypes. However, it is important to improve our understanding on the floral biology and reproductive system for this species to create new and efficient varieties.

Key words: Genetic variability, neglected plant, morphotype, crop phenology, growth parameters.


 INTRODUCTION

Fonio (Digitaria spp.) is one of the neglected and underutilized crops of West Africa. It is grown in  an  area stretching from Senegal to Lake Chad (Cruz et al., 2011). Well  adapted  to  local   pedo-climatic   conditions,   fonio resists drought due to its C4 metabolism and contributes to the maintenance of the environment by ensuring vegetal cover on ecologically sensitive and undervalued soils (Vall et al., 2008; Cruz et al., 2011). Poor in gluten, fonio is indicated for diabetics and those suffering from overweight and breastfeeding women (Vodouhe and Achigan Dako, 2006). It is rich in methionine and cysteine, two essential amino acids for humans and deficient in wheat, rice, maize and sorghum (Vietmeyer et al., 1996; Ballogoun, 2013).

In Benin, unlike other cereals such as maize, sorghum, millet and rice, that are cultivated everywhere throughout the country, fonio appears as an essentially local or endemic crop to the Atacora department in the northwest of Benin (Vodouhe et al., 2003). This crop has a socio-cultural importance for the Otamari ethnic group in Boukoumbé, the main producer community providing 74% of the national production (Dramé and Cruz, 2002; Ballogoun, 2013; Paraïso et al., 2013). It also plays an important role in food security in the population especially during the lean season when early varieties are used to curb famine.

Despite its potentials, fonio remains a marginal plant and long neglected in national research programs. Most of the research has put emphasis on the management of harvest and post-harvest. Little knowledge is available on varietal breeding  of  this  crop.  The  crop  is  thus  poorly known as far as the morphological, agricultural and biological characteristics are concerned. Today, it is hard to determine with accuracy, the varieties of fonio that are grown in Benin. The objective of this study was to assess and structure the morphological and agronomic variability of fonio ecotypes cultivated in Benin in order to improve their performance. 


 MATERIALS AND METHODS

The trial was conducted in 2015 at the experimental farm of the Faculty of Agronomy, the University of Parakou (9° 18' 57" North latitude, 2° 42' 5" East longitude, 362 m of altitude). The soil is poor in organic matter; it is of tropical ferruginous type and consists of about 22.40% of clay and silt, 1.43% of total carbon and 0.167% total nitrogen, or a C/N ratio of 8.56 (Azontondé et al., 2009).

Rains were regular and well distributed during the period of the trial (Figure 1). The wettest months were July, August, September and October. The total annual rainfall obtained in 56 days was 869 mm. The average daily temperature varied between 20 and 25 °C with a daily average of 22°C over the period of the study.

 

 

Twenty (20) accessions of fonio collected in the commune of Boukoumbé were evaluated in a randomized complete blocks design (RCBD) with three replicates. The experimental units consisted of 3 lines of 4 m each. Sowing was done at a spacing of 0.20 x 0.20 m (20 holes/line) with a pinch of seeds. The plants were reduced to one per hole 25th day after sowing. The alley between two consecutive experimental units was 0.4 m. The plots were kept clean by regular manual weeding until harvest. The  trial  block  was fenced with wood and protected with mosquito nets so as to exclude rodents and domestic animals. No fertilizer and pesticide treatments were applied. Harvesting was performed by mowing stems using sharp scissors for cutting the panicles. The plants were characterized using a descriptor adapted from the one utilized for the description of rice (Tia and Iliath, 2014). In total, 18 qualitative variables and 21 quantitative variables were analyzed so as to describe and partition the varieties in morphological groups. Qualitative variables were observed at the plot level. They consisted of visual observation of the coloration of different organs (stem, node, internode, leaf and panicle) during the vegetation, the flowering and the maturation phases (Table 1). Quantitative variables were measured, some at the plot level and the others at the plant level. These were collected from ten plants randomly chosen on the central lines of the elementary plots. The measures were phenology, plant growth and plant production parameters (Table 2).

 

 

 

R software version 3.1.3 was used for data analysis. Qualitative variables were studied by calculating, for each of them, the proportions of the accessions belonging to each category. The variability of quantitative traits was investigated through an analysis of variance. After an analysis of the correlation matrix, quantitative variables not correlated significantly (p>0.05) and differentiated the accessions as well as qualitative variables differentiated the accessions were submitted to a factor analysis of mixed data (FAMD) to identify morphotypes. The identified morphotypes were eventually described and the main differentiating traits for these morphotypes were determined by using Chi-squared test for categorical variables and λ-Wilk test for quantitative traits. For the later, the Tukey test was used to compare means when differences between morphotypes were significant.


 RESULTS

Qualitative description of accessions

Among the 18 evaluated variables, a dozen enabled distinctions between accessions. Some of the collars were purple green while others were purple. Also, some of the foliar limbs, internodes and stem nodes showed anthocyanin coloration while others were devoid of it. The panicles, whether horizontal or open, were either brown or red at maturity. The stems had an open port, semi-erected, or very open. There were also differences in the color of the plant base, the foliar limb, the ligula and the panicle exsertion between the accessions (Table 3).

 

 

Variability of morphology measurable characters

Differences in the  accessions  were  only  observed with plant height and panicle length (P < 0.01). Plant height, either in centimeters or number of nodes varied from simple to more than double. The shortest accession was AS1 and the longest was AS5. The same trend was observed for the height and node of panicle insertion. The shortest panicles were noted on AS12 and the longest on AS5. For the remaining variables, differences among accessions were not significant (Table 4). The analysis of the correlation matrix showed significant associations between some of these morphological variables (Table 5).Plant height (HP) was significantly (p<0.05) correlated with the width of the leaf under the panicle leaf (LargF), the height of panicle insertion (HIP), the node of panicle insertion (NIP), the number of nodes on the main stem (NN_Plt) and the number of racemes per panicle (rac_plt).

Width of panicle leaf (LargFP) was significantly (p<0.05) correlated with the width of the leaf under the panicle leaf (LargF), the length of the leaf under the panicle leaf (LongF) and the length of panicle leaf (LongFP). Panicle length (Lg_pan), raceme length (Lg_rac) and the number of  panicles  per plant  (Pan_plt) were not correlated with any other variables. For descriptions of morphotypes, five variables were retained: length of panicle leaf (longFP), plant height (HP), panicles length (Lg.pan), raceme length (Lg.rac) and number of panicles per plant (Pan.plt). These variables were considered sufficient to account for the accessions of morphometric variability since each of the other variables was correlated with either of these five.

 

 

 

Variability of agronomic characterization parameters

Despite  the  large  variations  observed  in  the   date   of emergence (4 to 10 days after sowing), number of tillers per plant (15 to 37), number of leaves per plant (51 to 134), and the harvest index (9 to 36%), there was no significant difference among accessions (p>0.05) for these parameters (Table 6). Nevertheless, there were significant variations among accessions for other parameters such as flowering and maturation dates. The early matured varieties were AS1 and AS8, whereas AS15 and AS10 matured late. 

 

 

Grain and biomass yields also showed significant variations between accessions. AS2 was the most productive accession, with more than 2 t/ha of grain yield, which  was  almost  nine  times  the yield of the least productive, AS1 (p <0.05). Significant differences (p <0.01) were observed among accessions for the biomass yield as well. AS1 recorded the lowest biomass yield, while AS5 recorded the highest (Table 6).

Pearson correlation matrix showed that the grain yield (Rdt_grain) was little correlated with the other variables; it was only correlated with the biomass yield (Rdt_biomasse). The biomass was correlated with most of the other variables except for the date of emergence (50% level), the number of leaves (Nb_feuilles) and the number of tillers (Nb_talle). There was also a strong correlation (p<0.01) between the date of flowering (50% CSE) and date of maturity (50% CSM), and between the number of leaves and number of tillers (Table 7).

 

 

For the identification of morphotypes, the date of emergence (50% level), date of flowering (50% CSE), harvest index (IR), grain yield (Rdt_grain) and number of tillers (Nb_talles) proved to be sufficient to describe agronomic variability of the studied accessions, given that each of the other  variables  was  correlated  with  one  or more of these selected parameters.

Regrouping of accessions in morphotypes

The categorical variables that distinguished accessions and quantitative parameters selected from the analysis of the correlation matrix were submitted to a factor analysis of mixed data (FAMD).

The first five axes explained 73.2% of the total information (cumulated variance) with all Eigen values ​​greater than 1. The first two axes explained 43.37% of the total variability (Table 8). The first axis was correlated with plant height (HP), earliness (50% CSE and 50% levée) and grain yield (Rdt_grain). It is mostly an axis characterizing vigor and earliness. It also describes the coloration of different parts of the plant, in particular, anthocyanin coloration of the foliar limb (ColAnt.LF) and, to a lesser extent, coloration of the internodes of the stem (ColAntEnNd),    racemes    (coul.racème)     and     ligula (Coul.lg) (Figure 2). The second axis was correlated with the number of tillers (nb_talles), the panicles length (Lg.pan) and grain yield (Rdt_grain). Regarding qualitative characters, it describes the color of the base of the plant (Coul.bas.pl), the port of the stem (Port.Tige) and panicles form (Exs.Pan and Type.Pan) (Figure 2).

 

 

 

The projection of accessions in the factorial plane formed by the first two axes allowed separation of the accessions into four groups (Figure 3). Axis 1 opposed the accessions of group 1, earlier, of short height and longest panicle leaves to those of group 4, later and more slender. Regarding qualitative traits, this axis also opposed accessions devoid of anthocyanin coloration at foliar limb and internodes (Group 1) to others which displayed anthocyanin coloration (Group 4).

 

 

The second axis opposed the accessions of group 2 and group 3, respectively located on the upper and lower side of this axis (Figure 3). Accessions of group 2 have a higher tillering aptitude, longer panicles and higher yields than accessions of group 3. Regarding qualitative traits, this axis opposed accessions equipped with horizontal panicles and very good exsertion (Group 2) to those equipped with open panicles and good exsertion (Group 3).

Identification of discriminating traits

Chi-square tests (χ²) showed that eight qualitative traits are significantly related to the classification of the accessions in morphological groups (Table 9). λ-Wilk test revealed that five (5) quantitative characters allowed discrimination of the identified morphotypes. These were the date of flowering (50%CSE), plant height (HP), length of panicle leaf (LongFP), raceme length (Lg.rac) and grain yield (Rdt_grain) (Table 10).

 

 

 

 


 DISCUSSION

The morphological and agronomic characterization of a crop is an important step in the management of genetic diversity (Manzano et al., 2001; Yobi et al., 2002; Radhouane, 2004). It is also a prerequisite towards the selection of improved varieties (Smith et al., 1991; Fraleigh, 1987). This work had helped to characterize for the first time in station in Benin, the diversity of ecotypes of fonio grown in Boukoumbé, one of the regions described as one of the areas of origin of this crop (Adoukonou-Sagbadja et al., 2006, 2007). The results showed significant differences between several morpho-phenological and agronomic characters. This demonstrates a high variability between accessions, which could be explained by the farmers’ practices of seed management coupled with the proximity to the study area with Togo, another region of origin for fonio where genetic diversity was described as very high (Adoukonou-Sagbadja et al., 2007). Indeed, several authors have shown that farmers' practices of seed management, including the exchange of varieties among farmers, are the source of an important diversity among populations of cultivated plants (Mckeye et al., 2001; Delaunay et al., 2008; Missihoun et al., 2012). The exchange of varieties among farmers, which has been described as the main mode of access to fonio seeds in this region (Sekloka et al., 2015), has been able to contribute to a significant increase in the diversity of ecotypes of fonio grown in Boukoumbé. This substantial phenotypic variability  could be an expression of a strong genotypic heterogeneity. Such morphological and phenological dissimilarities between accessions are often generated and maintained by diverse evolutionary processes. Agroecosystems are likely to exert widely varying selective pressures on genotypes (Sadiki and Jarvis, 2005). This is also the case with anthropic pressures (Robert et al., 2004). Indeed,the way seed is managed by farmers such as selective sorting, post-harvest technologies and agricultural practices lead to a selection involving the maintenance, and even the creation of a remarkable phenotypic diversity (Robert et al., 2005).

Our results revealed that the studied collection encompasses extra-early accessions of less than 90 days and late accessions of more than 100 days. These results consolidate and complement those found by Vodouhe and Achigan Dako (2003) which identified in Benin short cycle varieties (90 days on average) and long cycle varieties (about 120 days). Similar results regarding earliness of Fonio were also found in Niger by Saidou et al. (2014). Identifying early accessions is of great agronomic importance for varietal breeding of fonio in the current context where climatic variations are becoming recurrent. However, the floral biology of fonio remains poorly known and the mode of reproduction, preferentially autogamic for some varieties (Cissé, 1975), allogamic for others (Vodouhe and Achigan Dako, 2006) and essentially apomictic for Adoukonou-Sagbadja et al. (2010), has not make unanimity within the scientific community yet (Cruz et al., 2011). A good understanding of its floral biology and possibilities of achieving chromosomal mixings will enable to exploit this variability better and create more efficient varieties.

In this study, four fonio morphotypes were identified, whereas Dansi et al. (2010) found five morphotypes in Boukoumbé and surrounding areas. Indeed, Dansi et al. (2010) had done their ranking based on the results of field surveys and not experimentation in station, which already made a fundamental difference in the methodological approach used for the evaluation of accessions. Also, their work had covered the entire region of Atacora where fonio is grown in Benin and not only the main production commune of Boukoumbé which was the focus of this study. These reasons could be the differences in the number of morphotypes identified. Similar to these findings, Saidou et al. (2014) identified four morphological groups of fonio in Niger for the species Digitaria exilis through an experimentation of fonio accessions in station.

The results of this study suggest that the accessions grown in Boukoumbé were rather of D. exilis and not of Digitalia iburua. Indeed, the descriptions obtained in this study are similar to those found by Cissé (1975) for the species D. exilis. Moreover, referring to the work of Portères (1955), the leaves of D. iburua have long lashes near the basis, leaving behind the ligula which is membranous, round, wide, long of 1 to 2 mm, with a terminal panicle consisting of 4-10 sub-racemes. These results differ from this description and suggest that our accessions were rather of D. exilis. The factor analyses made it possible to identify a set of quantitative and qualitative characters that discriminate the accessions. Among these traits, descriptors of earliness, vegetative development and grain productivity ranked high.

Examples of these descriptors were the date of flowering, plant height, length of foliar limb and racemes length. The importance of these types of characters in structuring the diversity of vegetal populations has been demonstrated on maize (Moreno et al., 2006; N'da et al. 2014), cotton (Sekloka et al., in review), sorghum (Koffi et al., 2011) and many other crops. Although subject to environmental variations, these parameters should not be neglected by plant genetic resources managers since they have always been important in farmers’ environment and constitute important criteria for mass selection.


 CONCLUSIONS AND SUGGESTIONS

This study on agro-morphological characterization showed a significant diversity of ecotypes of fonio cultivated in Benin. They are diverse in their port, the color of their raceme, the presence or absence of anthocyanin at different levels of the stems and leaves. The earliest accessions had a sowing-maturity cycle of about three months, but grain yields were mostly less than or equal to 800 kg/ha. The most productive accessions recorded more than 1.5 t/ha of grain yield, with more than 2 t/ha for AS2. These accessions can be separated into four distinct morphological groups based on discriminating traits. The most discriminating qualitative traits were the color of collar, the intensity of green coloration of foliar limb; anthocyanin coloration and its distribution across different aerial organs of the plant, and the type and exsertion of panicles. As for quantitative parameters, the most discriminating were the date of flowering, plant height, length of foliar limb and racemes length. The morphological, phenological and agronomic variability demonstrated is sufficient to implement a program of varietal breeding based on varietal homogenization followed by multiple sites evaluation of the improved ecotypes. To sum up, these results contribute to a better knowledge of fonio ecotypes and therefore, allow a better management of the species variability for the benefit of the producers. However, it is important to improve understanding onthe floral biology and reproductive system of fonio in order to be able to achieve genetic mixings leading to new genotypes.


 CONFLICT OF INTERESTS

The authors have not declared any conflict of interests.



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