See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/320249590
Harnessing genetic resources and progress in plant genomics for fonio
(Digitaria spp.) improvement
Article  in  Genetic Resources and Crop Evolution · October 2017
DOI: 10.1007/s10722-017-0565-6
CITATIONS READS
0 171
5 authors, including:
Mathieu A.T. Ayenan Frejus SODEDJI
University of Ghana University of Abomey-Calavi
14 PUBLICATIONS   19 CITATIONS    7 PUBLICATIONS   1 CITATION   
SEE PROFILE SEE PROFILE
Charles Ikenna Nwankwo Katina Olodo
University of Hohenheim Institute of Research for Development
3 PUBLICATIONS   1 CITATION    2 PUBLICATIONS   0 CITATIONS   
SEE PROFILE SEE PROFILE
Some of the authors of this publication are also working on these related projects:
Seedball project View project
Matching grain quality attributes to the requirements of soybean processors in Benin View project
All content following this page was uploaded by Mathieu A.T. Ayenan on 01 May 2019.
The user has requested enhancement of the downloaded file.
Genet Resour Crop Evol
DOI 10.1007/s10722-017-0565-6
RESEARCH ARTICLE
Harnessing genetic resources and progress in plant genomics
for fonio (Digitaria spp.) improvement
Mathieu Anatole Tele Ayenan . Kpedetin Ariel Frejus Sodedji . Charles Ikenna Nwankwo .
Katina Floride Olodo . Mahulé Elysé Boris Alladassi
Received: 20 April 2017 / Accepted: 19 September 2017
 Springer Science+Business Media B.V. 2017
Abstract Fonio plays an important role in food crop notwithstanding its high nutritional and market
security and in income generation, in drought prone values. Breeding programs on fonio should first focus
areas of West Africa. This review aimed at highlight- on solving these problems to improve overall yield.
ing fonio production constraints, breeding objectives We explored the potential of mutation breeding,
and available genetic resources for fonio improve- somaclonal variation, somatic hybridization, molecu-
ment. Lack of institutional support to research, lack of lar markers, comparative genomics, individualized
improved seeds, parasitic weed infestation, grain targeting induced local lesions in genomes (iTIL-
shattering, lodging and toilsome of the post-harvest LING), genotype by sequencing and genomic selec-
handling are the major constraints limiting fonio tion to develop improved fonio varieties.
production. These factors lead to negligence of the Determination of ploidy level, wide collection and
M. A. T. Ayenan K. F. Olodo
West Africa Centre for Crop Improvement, College of UMR DIADE, Institut de Recherche pour le
Basic and Applied Sciences, University of Ghana, Développement, Université de Montpellier II,
PMB LG 30, Legon, Ghana 34394 Montpellier, France
K. A. F. Sodedji K. F. Olodo
Laboratory of Applied Ecology, University of Abomey- Laboratoire Mixte International Adaptation des Plantes et
Calavi, 01 BP 526, Cotonou, Benin microorganismes associés aux Stress Environnementaux
(LMI LAPSE), Centre de recherche de Bel-Air, Dakar,
K. A. F. Sodedji  M. E. B. Alladassi (&) Sénégal
College of Agricultural and Environmental Sciences,
Makerere University, P.O. Box 7062, Kampala, Uganda K. F. Olodo
e-mail: alladassi.meb@gmail.com Centre d’Etudes Régional pour l’Amélioration de
l’Adaptation à la sécheresse (CERAAS), BP 3320, Route
C. I. Nwankwo de Khombole, Thiès, Sénégal
Institute of Soil Science and Land Evaluation, University
of Hohenheim, Emil-Wolff Str. 27, 70599 Stuttgart, K. F. Olodo
Germany Université Cheikh Anta Diop, BP 5005, Dakar-Fann,
Dakar, Sénégal
K. F. Olodo
Laboratoire National de Recherche sur les Productions
Végétales (LNRPV/ISRA), BP 31200, Route des
hydrocarbures, Bel-Air, Dakar, Sénégal
123
Genet Resour Crop Evol
characterization of fonio genetic resources, definition Fonio plays a critical role in food security and income
of core reference set collection, and exploitation of generation in marginal areas in West Africa. It is still
heterosis are some key research areas that would be of referred to as either an old crop or women’s crop
great interest in fonio improvement. However, this can (Vodouhè et al. 2007; Vall et al. 2011; Small 2015).
only be achieved with adequate funding and institu- Fonio has several end products and consumption
tional support. forms (e.g., porridge, couscous, and paste for human
consumption) (Vodouhè et al. 2007). It has gained
Keywords Breeding  Digitaria spp.  Genetic interest in food industry (formulation of bread,
resources  Genomics  Gramineae  Neglected and cookies, sour dough, drinks, non-fermented steamed
underutilized crop and granulated dumpling products) resulting in
increased demand and market value of the crop
(Jideani and Jideani 2011). High traditional and
customary values are attached to fonio, which is
Introduction reserved for chieftaincies, royalties, special traditional
occasions and ancestral worship (Adoukonou-Sag-
By 2050, world’s population is expected to reach 9 badja et al. 2006; Blench 2012). The straw serves as
billion (Nelson et al. 2010; Godfray et al. 2012). This good livestock feed (Clottey et al. 2006; Vodouhè
will result in an exceedingly increased demand for et al. 2007).D. exilis andD. iburua contain on average
food, which is estimated to reach 100–110% in 2050 per 100 g of edible grains about 8% protein, 75%
compared to its level in 2005 (Tilman et al. 2011). carbohydrate and 6.8% dietary fiber (Table 1) (Brink
However, crop production has been negatively 2006a). Due to its higher content of essential amino
affected by climate change (Nelson et al. 2010), acids, methionine and cysteine compared to wheat,
competition for agricultural land and energy and water rye, barley, rice sorghum and maize, it is referred to as
for irrigation (Khan et al. 2006; Ronald 2011). These the most nutritious cereal grain (Table 1) (Vodouhè
phenomena will have severe negative effects on food et al. 2007; Koroch et al. 2013; Ballogou et al. 2014).
security (FAO 2016). In addition, high risks of The grains are gluten-free and suitable for curing
malnutrition arising from hidden hunger due to low coeliac disease (Taylor et al. 2006) and its consump-
diet food consumption might become significant tion is beneficial for lactating women and diabetic
(Kennedy et al. 2003; Bouis and Welch 2010; Ngozi patients (Vodouhè et al. 2007).
2013). With the current production model based on Investing in neglected crops has been considered as
few crops species, it may be challenging to meet the cost-ineffective regarding their local production and
global food demand. Sustainable intensification and less interest in international trade. However, owing to
diversification of agricultural and food systems fonio importance, there have been several calls for
through promotion of neglected crops species is being breeding for improved fonio varieties, and conserva-
encouraged (Kahane et al. 2013; Padulosi et al. 2013; tion and uses of its genetic resources (Barnaud et al.
Massawe et al. 2016). One of such crops is fonio, 2013). Additionally, calls for improved cultural prac-
which has high agricultural, nutritional and commer- tices, processing and marketability have been reported
cial values. (Gigou et al. 2009; Koreissi-Dembélé et al. 2013;
Fonio is one of the oldest domesticated cereals in Ballogou et al. 2014). There has been recent research
West Africa dating back as far as c. 2400 BC (Larson on fonio improvement and related cereal species that
et al. 2014). The earliest evidence of its cultivation need to be discussed to orient future research on the
dated to c. 1000 and 400 BC in Mali (Takezawa and crop.
Cisse 2004). It belongs to the Gramineae family, the In this review, we addressed the following ques-
subfamily Panicoideae and the genus Digitaria. Two tions: (1) what factors constrain fonio’s production?
major known cultivated fonio species are the white (2) what has been done to improve fonio? (3) what
fonio; Digitaria exilis (Kippist) Stapf and the black could be the objectives of fonio breeding programs?
fonio Digitaria iburua Stapf (Hilu et al. 1997; (4) what genomic resources and breeding techniques
Adoukonou-Sagbadja et al. 2006). The closest grass are available for use to achieve these objectives?
genera to Digitaria are Cenchrus and Setaria (Fig. 1).
123
Genet Resour Crop Evol
Fig. 1 Phylogenetic tree showing relationship among three (Letunic and Bork 2016) based on the NCBI taxonomy. D.
fonio species and other grasses. The tree was generated using iburua is not in the NCBI database and it could be included in
phyloT (http://phylot.biobyte.de) and visualized with (iTOL) v3 the tree
Table 1 Nutritional content per 100 g of fonio and other cereal crops. Source: Badu-Apraku and Fakorede (2006), Balole and
Legwaila (2006), Brink (2006a, b), Vodouhè and Achigan Dako (2006)
Species Protein Carbohydrate Dietary Tryptophan Methionine Phenylalanine Threonine Valine Leucine
(g) (g) fiber (mg) (mg) (mg) (mg) (mg) (mg)
D. exilis 7.1 74.4 7.4 111 441 402 315 457 772
D. iburua 8.9 75.6 6.2 215 355 803 389 614 1395
Rice 6.7 81.3 3.8 87 140 330 230 390 560
Sorghum 11.3 74.6 – 124 169 546 346 561 1491
Maize 9.4 74.3 7.3 67 197 463 354 477 1155
Wheat 15.4 68 12.2 195 230 724 433 679 1038
Methodology PubMed Central and JSTOR. No date coverage was
specified during the search. In each of the database,
Literature search was conducted in various databases key words related to the defined questions were used to
including Google Scholar, Web of Science, Scopus, conduct the literature search. Fonio diversity, fonio
123
Genet Resour Crop Evol
germplasm were used to collect literature on fonio Challenges in fonio production system and post-
diversity and germplasm. Fonio production system, harvest
fonio cultivation, fonio post-harvest and fonio seed
system were used to get published and unpublished Fonio farmers lack good agronomic practices and
(reports) works on fonio production systems, post- improved varieties, which are needed to reach the
harvest handling and challenges. Literature on avail- yield plateau. Farmers’ access to improved varieties
able genomic resources was obtained using key words and good-quality seed is of great importance to
including New Generation Sequencing, TILLING and increase productivity and production (Louwaars and
plant breeding, Genotyping by sequencing: applica- de Boef 2012). However, as in other minor crops, there
tion in plant breeding, Genomic selection, mutagen- is no formal seed system in fonio (Ahmed et al. 2009;
esis and plant breeding, micro-propagation and plant Ayenan and Ezin 2016). Designing of sustainable seed
breeding and genomic resources and neglected crops systems that are rooted in and led by the community
and genomic tools and cereals breeding. Papers’ title and can enable production and timely delivery of good
and/or abstract were screened and selected based on quality seeds to farmers is deemed important.
their relevance to the questions being investigated. A low and relatively constant yield of 0.2–0.9
-1
Additionally, the reference lists or bibliographies of t ha has been reported across fonio growing areas
the selected papers were checked manually for articles (Adoukonou-Sagbadja et al. 2006; Dachi and Gana
of interest. The selected papers were subjected to 2008; Adoukonou-sagbadja 2010). Even though the
critical analysis. crop is known to thrive well in marginal areas,
prolonged or terminal drought stress have also been
reported as important emerging constraint to fonio
Results and discussion production especially in semi-arid zones, character-
ized by unpredictable, irregular and shorter
Fonio production (2–3 months) rainy season (Dansi et al. 2010).
In Benin, farmers reported some parasites including
Fonio is cultivated across West African Savannah ‘‘insects’ larvae’’, which consumed limbs during
from Lake Chad to Senegal where it serves several drought period and insects called ‘‘fonio mosquitoes’’
millions of indigenous people as a staple and strategic which feed on spikelet (Adoukonou-sagbadja 2010).
crop for food security during lean periods (Barnaud Susceptibility of fonio to the fungi Phyllachora
and Billot 2011; Vall et al. 2011). Guinea is reported to sphearosperma and Helminthosporium spp. as well
be the top producer of all fonio growing countries with as to rust caused by Puccinia cahuensis were also
a cultivation area of about 459,563 ha and a produc- reported (Adoukonou-sagbadja 2010). This underlines
tion of 479,985 t in 2014, accounting for about 75% of the need for further study on the identification,
the total fonio production (FAOSTAT 2017). West incidence and severity of these pests in order to design
African farmers cultivate more than 550,000 ha of adequate management strategies. Weeds also con-
land area per annum (FAOSTAT 2017). Fonio tribute to some extent to the yield gap as fonio plants
production objective varied across growing areas. In are low weed competitors (Adoukonou-Sagbadja et al.
semi-arid areas of Burkina-Faso, Guinea and Mali, 2006). Striga aspera and Striga hermonthica are
fonio is mainly grown to supplement food shortage reported as the most important fonio parasitic weeds
during lean periods while in sub-humid regions of in West Africa (Hillocks et al. 1996; Gigou et al.
these countries, fonio is mainly grown for crop 2009). Other production challenges include shattering
diversification (Vall et al. 2011). The wide ecological and lodging. It has been estimated that shattering
adaptability of fonio gives it high potential as causes up to 30% of grain loss in fonio (Vodouhè et al.
promising crop in future to provide food security, 2007). While the production constraints are very
not only in West Africa where it is currently being important and efforts must focus on improving all
cultivated, but also in other parts of Africa where its productions aspects, there are aspects such as post-
cereal counterparts thrive (Small 2015). harvest challenges which should not be overlooked if
fonio production is to be promoted.
123
Genet Resour Crop Evol
Fonio grains are small in size (1000 grains weigh on Breeding for delayed shattering
average 0.44 g); their handling and processing (thresh-
ing, winnowing, dehulling, whitening) are hard and Breeding fonio for resistance to uncontrolled seed
toilsome task in absence of appropriate technologies. shattering is deemed important (Fig. 2). Loss of
These challengeshavebeen for longand still being some shattering habit was a major event in domestication
of the reasons behind the neglected status of this crop process of cereals (Konishi et al. 2006; Lin et al.
(Ballogou et al. 2014). However, progress has been 2012). Seed shattering in fonio is believed to be both
made across some of the fonio growing areas (Mali, environmentally and developmentally controlled (Pat-
Burkina-Faso, Guinea) with the designing of machines terson et al. 2016). Using comparative genomics, the
to reduce the tiresomeness of post-harvest operations ortholog of the gene qSH1, a shattering gene in rice
(Marouzé et al. 2008). According to the authors, this has and members of the Agamous-related family of genes
led to an increasing economic potential of the crop. such as JOINTLESS 1–3, SH2, and SH3, the free-
There is still, however, a gap to be filled at both national threshing locus Q of wheat, as well as the Arabidopsis
and regional levels as far as the availability and the abscission-associated gene NEVERSHED (Patterson
affordability of the threshing machines are concerned. et al. 2016) were identified in fonio. In several
cultivated cereals, loss of shattering was caused by
Potential breeding objectives in fonio few mutations at major QTLs controlling the trait.
Thus, SNP mutation at qSH1 locus in Oryza sativa
Actors (farmers, processors, consumers, etc.) in the (Konishi et al. 2006), three mutations in Shattering1
value chains of a particular crop have specific traits of (Sh1) locus in Sorghum (Lin et al. 2012) and SNP
interest and needs upon which they select their mutation at GL4 locus in Oryza glaberrima (Steud.)
preferred varieties. In regards to the preferred traits (Wu et al. 2017) were responsible for loss of shattering
in a specific fonio variety, Dansi et al. (2010) reported in these crops. These mutations led to loss-of-function
eight preference criteria namely extra early-maturity, of genes underlying seed shattering. Further research
large grain size, culinary characteristics, ease of to map QTLs controlling seed shattering in fonio and
processing, productivity, ease of harvesting, grain understanding of the molecular and physiological
size, storability and drought tolerance (Fig. 2). These basis of this trait are important to enable breeders to
criteria must guide the implementation of fonio select most appropriate methods to develop resistant
breeding programs. varieties to uncontrolled seed shattering. When few
major QTLs underlie grain shattering in fonio like in
other cereals, use of mutation breeding could be
efficient in developing non-shattering varieties that
Fig. 2 From breeding
objectives to outcomes for
1 Reduced Resistance 
Grain size (ease post- Organoleptic traits Extra early 
farmers and consumers (1, 2 shattering  to lodging harvest handling, (texture, cohesiveness, 
and 3 = breeding increase weight) aroma, etc) 
maturing 
objectives; 4 = outcomes
for farmers and consumers)
2 Increased Yield  
3 Breeding for dual purpose 
(grain and fodder) 
Meeting consumers’ 
4 Increased Animal Increased resilience: preferences: 
production feeding to climate change and Increased market value cropping systems 
123
Genet Resour Crop Evol
would result in overall grain yield improvement. This for the genes TaGS5-3A (Ma et al. 2016) and TaGW2
approach was successfully used to develop non- (Su et al. 2011) and in maize for the genes ZmGW2-
shattering forage rice variety (Kato et al. 2006). CHR4 and ZmGW2-CHR5 (Li et al. 2010). Functional
genomics will then come in handy, to determine if
Breeding for resistance to lodging these homologs have similar function in fonio as in
rice, wheat and maize. Finally, through genetic
Lodging poses limitations to fonio yield but till date, engineering, a significant increase in fonio grain size
no breeding effort has tackled this issue (Fig. 2). Wide could be achieved through the development of GW2-
germplasm screening for both cultivated and wild underexpressing lines. Indeed, increased grain size
relatives could help to identify materials resistant to and weight have been obtained by Song et al. (2007) in
lodging which could be used to develop improved rice after creating an antisense mutation that results in
varieties with stronger culms and/or with dwarfism a truncation of 310 amino acids and by Hong et al.
genes. Additionally, two alternative options, involving (2014) in wheat after transforming a TaGW2-RNA
Gibberellic Acid (GA) biosynthesis pathways which interference cassette in the variety Shi4185 with small
has been extensively used in other cereal crops (rice grain.
and wheat), could potentially be exploited to develop
lodging resistant fonio varieties (Peng et al. 1999; Breeding for high-yielding varieties
Sasaki et al. 2002; Asano et al. 2009; Okuno et al.
2014). The first option is the creation of GA deficient Developing farmers’ preferred and high yielding
mutants to generate dwarf lines which will be further varieties is of utmost importance to upgrade the status
evaluated for agronomic performance. The second of the crop. Although some phenotypic evaluations for
option, which is now being explored in rice breeding, high-yielding genotypes have been recently con-
targets not only the breeding for resistance to lodging ducted, no major breeding efforts were made for the
but also the production of high biomass by creating genetic improvement of fonio’s productivity. How-
mutants with over-production of GA. This strategy ever, from the multi-locational participatory variety
enables formation of larger culm diameters and/or selection implemented in Mali, seven landraces were
lignin accumulation resulting in improving the break- identified as earlier-maturing and higher yielding than
ing-type lodging resistance (Okuno et al. 2014). the local checks with the landrace ‘‘Fini of Samakou
Dansira-1’’ which had a stable yield of 1.4 t ha-1
Breeding for larger grain size (Sanogo and Hash 2014). This indicates that signifi-
cant improvement could be made through the collec-
Grain size is one of the two determinants of grain tion and evaluation of germplasm at regional level.
weight which is a major component of yield, for most Besides, grain yield being a quantitative trait, function
cereal crops (Brocklehurst 1977). In fonio, small grain of several yield component traits, breeding for resis-
size has been, for long, a key limiting factor for its tance to shattering and lodging and increased grain
processing and low grain-yield. Although to date, no size result into an improvement of both fonio’s
breeding effort has really addressed this challenge, the production and productivity (Fig. 2).
significant progress made in other cereals like rice and Fonio’s straw is good livestock feed for both
wheat could be advantageous. The use of comparative monogastric (Clottey et al. 2006) and ruminant
genomics is a potential avenue for breeding for large animals (Vodouhè et al. 2007). Breeding varieties
grain size in fonio. Two genes OsGS5 and OsGW2 with high biomass yield can be useful for animal feed
were identified in rice as grain size and weight especially in semi-arid zones characterized by unpre-
regulators; while OsGS5 encodes for large grain size dictable rainfall leading to low availability of fodder
by increasing the number of cells generated during cell and pastures. Furthermore, fonio is well appreciated in
division (Li et al. 2011), OsGW2 affects negatively the some sub-Sahara African countries which are at high
grain size and weight (Song et al. 2007). The risk of micronutrient deficiency, thus an improvement
homologous genes of OsGS5 and OsGW2 could of fonio for high density micronutrient characteristics
therefore be cloned and mapped on fonio genome will help to address the food security and nutrition
through comparative genomics as it was done in wheat challenges in this area.
123
Genet Resour Crop Evol
Germplasm and conservation problematic: kept in international genebanks to limit the risk of their
an incomplete and unstandardized collection loss.
There is no collection of wild fonio and black fonio
Plant breeders heavily rely on genetic variability at regional level; and this may limit the extent of
without which little or no progress can be achieved. exploiting the secondary and tertiary gene pool of
Readily available and well characterized germplasm fonio (Olodo 2016). For D. exilis, it is essential to
collections are important for breeders. D. exilis has a extend collection efforts in other countries where
wide distribution from Senegal to Lake Chad while D. presence of fonio is reported in the literature [such as
iburua is mainly encountered in Nigeria, Benin and Gambia, Côte d’Ivoire, Guinea Bissau (FAOSTAT
Togo (Adoukonou-sagbadja et al. 2004; Blench 2012). 2017), Sierra Leone (National Research Council
Based on this geographical distribution, important 1996), Dominican Republic (Morales-payán et al.
efforts have been made to collect accessions of D. 2002)] (Fig. 3). It is worth noting that while fonio
exilis. The first regional mission of fonio genetic germplasm might have been collected and kept in
resources collection was undertaken by Clément and national genebanks in some of these countries, to the
Leblanc (1984) in six countries (Benin, Niger, Togo, best of our knowledge there is no record of their
Mali, Burkina-Faso, Guinea) where they collected 641 existence. This emphasizes the importance of keeping
accessions of D. exilis which are maintained by IRD duplicates of genetic resources in international
(Institut de Recherche pour le Développement) in genebanks not only to ease access to these materials
Montpellier (Clément and Leblanc 1984). Germplasm to potential users but also to avoid their loss. Threat on
collection missions are also conducted at national genetic erosion is reported in some growing areas (e.g.
level (e.g. Mali, Guinea, Nigeria and Benin) for Togo) (Adoukonou-sagbadja et al. 2004), hence
genetic diversity studies and nutritional characteriza- indicating the importance of developing strategies to
tion. Duplicates of some of these fonio accessions are conserve its diversity. We propose a gap analysis
Fig. 3 Distribution of Digitaria spp. crops in West Africa with Biologiques Tropicales de Montpellier (CRB-T)) were reported
countries where from accessions kept in genebanks (U.S. or not. Source: Adapted from Blench (2012)
National Plant Germplasm System and Centre de Ressources
123
Genet Resour Crop Evol
(Michael et al. 1993) to identify the areas where iburua and the wild relatives (D. longiflora and D.
accessions are not collected or are under-represented ternata) can be assessed for more comprehensive
in genebanks. This will require collaboration among diversity analysis. Phylogenetic studies in fonio in
various institutions involved in maintaining fonio order to establish domestication and distribution are
germplasm. Doing so can help gather basic informa- also made possible with these markers.
tion (geographical coordinate, species, date of collec-
tion, number of available accessions) on fonio Fonio reproductive system
germplasm. After a wide collection, morphological,
molecular and nutritional characterization of the Fonio mating system was believed to be inbreeding or
germplasm should be carried out to develop a core apomictic (Adoukonou-sagbadja et al. 2010). Recent
reference set and make efficient use of the genetic evidence using micro-satellite markers revealed that
resources (Glaszmann et al. 2010). That will enhance the crop is predominantly self-fertilized with an
germplasm exchange and help breeders in selection of outcrossing that could be up to 1.7% (Barnaud et al.
desired materials for their breeding programs. 2017). Further investigation could address the poten-
tial to exploit heterosis in fonio to develop hybrid
Genetic diversity and available genomic resources varieties. However, up to now no successful experi-
ence in artificial hybridization has been reported.
The analysis of nuclear DNA contents using flow Traditional hybridization is very-difficult in fonio as
cytometric confirmed the close relationship between hand emasculation is often unsuccessful due to the
the cultivated fonio D. exilis and D. iburua and the extraordinarily miniature nature of its floral organs
wild species Digitaria longiflora (Retz.) Pers. and (Vodouhè and AchiganDako 2006). Research avenues
Digitaria ternata (A. Rich.) Stapf (Adoukonou-Sag- in somatic hybridization can be explored. This tech-
badja et al. 2007) as reported earlier by Hilu et al. nique circumvents sexual reproduction and helps
(1997). D. longiflora has been reported to have overcome crossing barriers (Jansky 2006). Thus, intra
interesting agronomic traits, including erect habit, and interspecies crossings can be envisioned in fonio.
strong and resistant to lodging; long panicle with high Nevertheless, this will require the development of
number of grains and large seeds (Dansi et al. 2010). protoplast fusion, hybrid cell selection and hybrid
Morphological characterization of 67 white fonio regeneration methods.
accessions collected in Niger showed high variability
in both qualitative (color of the grains and stems) and Available genomic resources and breeding
quantitative traits (yield, dry biomass, internode techniques for fonio
length and growth cycle) (Saidou et al. 2014).
Few genetic diversity studies have been done on Next-generation sequencing (NGS) provides research-
fonio based on molecular markers. Hilu et al. (1997) ers with cost effective and faster methods for gener-
and Kuta et al. (2003) assessed the genetic diversity in ating large quantities of sequence data (Mardis 2008;
10 fonio accessions collected from Togo and Nigeria, Varshney et al. 2009; Egan et al. 2012). Interestingly,
respectively. Adoukonou-Sagbadja et al. (2007) used genome synteny and the increasing availability of
Amplified Fragment Length Polymorphisms (AFLPs) Plant Genome Databases (Phytozome, Ensembl
to assess the genetic diversity among 122 accessions Plants, PlantGDB Genome Browser, National Center
of D. exilis and D. iburua collected from Benin, for Biotechnology Information, etc.) have made
Burkina Faso, Guinea, Mali and Togo. The AFLPs possible comparative genomics. Thus, genomic
markers clearly separated D. exilis from D. iburua. resources developed in other well studied cereals with
Low genetic diversity was found inD. iburuawhileD. available genome sequences could be exploited in
exilis showed moderate genetic diversity. A set of 38 fonio (Barnaud et al. 2013; Saha et al. 2016).
nuclear microsatellite markers (SSR) were developed Increasing availability of genome sequence in cereals
of which 21 exhibited polymorphism across a range of including Zea mays L., Triticum aestivum L., Oryza
36 accessions of D. exilis collected from Benin, sativa L., Sorghum bicolor (L.) Moench, Setaria
Burkina Faso, Togo, Guinea and Mali (Barnaud et al. italica (L.) P. Beauv. could therefore be harnessed for
2012). Transferability of these SSR markers to D.
123
Genet Resour Crop Evol
Fig. 4 Integrated tools and
TECHNIQUES FUNCTIONS 
techniques for fonio
improvement
Somaclonal variation 
iTILLING Germplasm Generate mutants 
enrichment 
MutMap+ 
Comparative genomic 
(Available genome in Gene discovery, 
related species) Gene function,  
QTL identification, Development 
Plastomic Molecular markers of improved 
discovery 
-Germplasm varieties 
collection Genotyping by 
-Diversity Sequencing
analysis 
-Definition Genomic Selection Estimation of marker 
of reference effects and prediction 
core set of Genomic breeding 
Gap Analysis values 
gene discovery, mapping of molecular markers, and Recent progress in genomic and breeding methods
gene annotation in fonio. could help to circumvent the limited knowledge on
Rapid decrease in costs of DNA sequencing fonio’s genome, leading to significant genetic gain
combined with advances in bioinformatics allowed (Fig. 4). Genomic Selection (GS) is becoming popular
the use of genotyping-by-sequencing (GBS), DNA in plant breeding programs and relies extensively on
sequencing (RAD-seq) and Diversity Arrays Technol- the use of molecular markers. Most of the variation
ogy (DArT) for SNP discovery and genetic analysis due to minor QTL is captured in the statistical
even in non-model crops (Elshire et al. 2011; Saha prediction model (Meuwissen et al. 2001). In the
et al. 2016). These genomic tools have a wide range of absence of data on genome sequence and genetic maps
applications, including but not limited to genetic in fonio, using GS with SNP mined from genotyping
diversity study, molecular markers discovery, genome by sequencing tags could yield quick results in traits
mapping, whole genome sequencing and gene discov- improvement.
ery (Varshney et al. 2009; He et al. 2014). In fonio, Research on fonio plastome seems to gain interest
GBS can be used to mine SNPs for population in recent studies, namely identification of chloroplast
structure and diversity studies and to identify mar- SNP (Scarcelli et al. 2011) and a snapshot of
ker-trait associations (Fig. 4). transcriptome fonio Niatia seedling (Sarah et al.
Molecular marker-based genetic linkage maps or 2017). In addition, a cost-effective protocol has been
genome-wide association studies (GWAS) important developed to sequence complete chloroplast genome
for detection of quantitative trait loci (QTLs) agro- for several individuals at a single MiSeq run and this
nomic traits (yield, grain size, resistance to lodging protocol has been validated with success in fonio
and shattering, etc.) are yet to be carried out in fonio. (Mariac et al. 2014). The availability of complete
Association of molecular markers to QTLs related to chloroplast genome sequence is an opportunity for
traits of interest could be established through compar- pre-breeding research (phylogenetic, domestication
ative genomics approach (Saha et al. 2016). This and genetic variability) in fonio (Mariac et al. 2014;
would pave way for the use of marker-assisted Daniell et al. 2016). Chloroplast genome has already
selection (MAS) in breeding for improved fonio been used in phylogenetic studies of commercially
cultivars. important crops (e.g.: rice, cotton) and has helped in
123
Genet Resour Crop Evol
efficient use and conservation of genetic resource 2003). The former would increase the frequency of
(Daniell et al. 2016). induced mutation while the latter would help to
Generating mutants can be a powerful approach to recover somaclones with desirable traits (Afrasiab
enrich diversity in fonio, carrying out basic research and Iqbal 2010). However, technical challenges
on gene function and subsequently identify materials related to selection of stable mutants should be
which can be advanced and released as varieties. Tools addressed through training of laboratory technicians
are now available to plant biologists to identify with and establishment/equipment of tissues culture facil-
precision individuals carrying mutation of interest. ities. In this regards, the assistance of the International
Targeting Induced Local Lesions IN Genomes (TIL- Atomic Energy Agency (IAEA) in Vienna (Austria)
LING) was initially developed in Arabidopsis to can be requested in materials irradiation and other
screen genes for mutations using a PCR-based assay technical issues in mutation breeding. Use of chemical
(McCallum et al. 2000). This approach has been mutagens can be explored as well.
widely adopted in many other plants species (Bush and
Krysan 2010). Modifications have been made on Institutional support for fonio research
TILLING leading to new procedures such as individ-
ualized TILLING (iTILLING), making mutation- Because of scarce resources devoted to fonio research,
screening more practicable with reduced investment we advocate that a research hub should be created for
(Bush and Krysan 2010). Applying iTILLING does fonio in West Africa. This approach will help to
not require M2 seeds to be catalogued; they can be efficiently use available resources by avoiding dupli-
harvested in bulk from M1 population, hence, consid- cation. The hub can be hosted by one of the national
erable time is saved (Bush and Krysan 2010). The research institutes where basic research on the crop as
iTILLING procedure would then be suitable for fonio well as varieties development will be done to cope
because of its tiny seeds (Patterson et al. 2016). with biotic and abiotic stresses and meet consumers’
Another approach for identifying mutation is preferences. In this regards, noticeable advance in
MutMap?, initially developed and applied on rice to collaboration among national research institutes in
detect causal mutations by comparing SNP frequen- West Africa and international research centers have
cies of bulked DNA of chemically-induced mutant and been made (Vodouhe et al. 1998; Barnaud and Billot
wild-type progeny of M3 generation derived from 2011). For example, through these collaborations,
selfing an M2 heterozygous individual (Fekih et al. protocols have been defined in order to standardize
2013). Because it does not require artificial crossing, collection of fonio genetic resources (see Arcad
which is a challenge in fonio, MutMap? has been Project, https://www.arcad-project.org/). Such a
identified as a potential technique for gene isolation regional approach has been identified as a key to the
and identification of genes controlling important success in pigeonpea breeding in Eastern and South
agronomic traits (Fekih et al. 2013). Africa (Kaoneka et al. 2016). Implement this approach
Tissue culture harbors interesting perspectives for to valorize fonio will require, as in other neglected
fonio improvement (Kuta et al. 2003). A protocol for crops, public funding (Stamp et al. 2012).
fonio tissue culture has been established (Ntui et al.
2010) and can be exploited to generate somaclonal
variation (Larkin and Scowcroft 1981). Somaclonal Conclusion
variation has been reported to be very successful in
crops with limited genetic systems (e.g., apomicts, Fonio has potential that can be tapped into as a
vegetative reproducers) and/or narrow genetic bases nutritious food security and commercial crop. How-
like fonio (Krishna et al. 2016). It was exploited to ever, low yield (grain and biomass), poor agronomic
develop a mutant SE7 with reduced plant height and practices, lack of improved varieties, difficulty in post-
increased grain yield in finger millet (Baer et al. 2007), harvest handling and processing are some of the
suggesting its potential use in fonio to enrich diversity factors that still hinder the exploitation of this
and develop improved variety. potential. Herein, research avenues for fonio’s
Combination of mutagenesis with micro-propaga- improvement through determination of ploidy level
tion could help in fonio improvement (Kuta et al. especially in the wild relatives, wide collection and
123
Genet Resour Crop Evol
characterization of fonio’s genetic resources, defini- agro-ecological zones of West Africa. Theor Appl Genet
tion of reference core collection and setting up of 115:917–931. doi:10.1007/s00122-007-0618-x
Adoukonou-sagbadja H, Wagner C, Ordon F, Friedt W (2010)
breeding programs to tackle these constraints are Reproductive system and molecular phylogenetic rela-
explored. Our evaluation showed that there are tionships of fonio millets (Digitaria spp., Poaceae) with
available techniques and genomic resources that can some polyploid wild relatives. Trop Plant Biol 3:240–251.
be leveraged on to collect, conserve and develop doi:10.1007/s12042-010-9063-0
Afrasiab H, Iqbal J (2010) In vitro techniques and mutagenesis
improved fonio varieties. In addition, strengthening for the genetic improvement of potato cvs. Desiree and
multi-stakeholder approaches and collaboration to Diamant. Pak J Bot 42:1629–1637
advance fonio research through public funding might Ahmed HMI, Gregg BR, Louwaars NP (2009) Seed systems for
play crucial roles at closing these research-call gaps in underutilized crops. Acta Hortic 2:459–464
Asano K, Hirano K, Ueguchi-Tanaka M, Angeles-Shim R et al
fonio. In this review, we clearly presented what has (2009) Isolation and characterization of dominant dwarf
been done so far with regard to fonio improvement, the mutants, Slr1-d, in rice. Mol Genet Genom 281:223–231
way forward. The information provided is valuable for Ayenan AMT, Ezin VA (2016) Potential of Kersting’ s
scientists interested in fonio research and offers a good groundnut [Macrotyloma geocarpum (Harms) Maréchal &
Baudet] and prospects for its promotion. Agric Food Secur
roadmap for sustainably harnessing genetic resources 5:1–9. doi:10.1186/s40066-016-0058-4
and research capacities to boost fonio production and Badu-Apraku B, Fakorede MAB (2006) Zea mays L. In:
utilization. PROTA (Plant Resources of Tropical Africa/Ressources
végétales l’Afrique tropicale)
Acknowledgements We are thankful to the West Africa Baer GI, Emets AI, Stadnichuk NA et al (2007) Somaclonal
Agricultural Productivity Program (WAAPP/PPAAO 2A variability as a source for creation of new varieties of finger
CERA 58 ID06) for funding research activities of KFO and millet (Eleusine coracana (L.) Gaertn.). Cytol Genet
Deutscher Akademischer Austausch Dienst (DAAD) for grants 41:204–208. doi:10.3103/S0095452707040020
awarded to KFO. We are grateful to Alcade C. Segnon for Ballogou VY, Soumanou MM, Toukourou F, Hounhouigan JD
designing the map and Dr. Moussa D. Sanogo of Institut (2014) Indigenous knowledge on landraces and fonio-
d’Economie Rurale (IER),Mali for his worthful discussions.We based food in benin. Ecol Food Nutr 53:390–409. doi:10.
also thank the two anonymous reviewers for their valuable 1080/03670244.2013.811388
comments. Our warmest gratitude goes to our respective Balole TV, Legwaila GM (2006) Sorghum bicolor (L.) Moench.
institutions for facilitating access to information during the In: PROTA (Plant Resources of Tropical Africa/Res-
course of this review. sources végétales l’Afrique tropicale)
Barnaud A, Billot C (2011) De la connaissance à la valorisation
du fonio. Cah Agric 20:310–312
Compliance with ethical standards
Barnaud A, Vignes H, Risterucci A-M et al (2012) Development
of nuclear microsatellite markers for the fonio, Digitaria
Conflict of interest The authors declare that they have no
exilis (Poaceae), an understudied West African cereal. Am
conflict of interest.
J Bot 99:e105–e107. doi:10.3732/ajb.1100423
Barnaud A, Vigouroux Y, Barry B et al (2013) From advanced to
underutilized crops: making fonio benefit from research on
References major cereals in Africa. In: Massawe F,Mayes S, Alderson
P (eds) International symposium on underutilized plants
species ‘‘crops for the future—beyond food security.’’ Acta
Adoukonou-sagbadja H (2010) Genetic characterization of tra-
ditional fonio millets (Digitaria exilis, D. iburua STAPF) Hortic (ISHS), vol 979. pp 421–430
Barnaud A, Vigouroux Y, Diallo MT et al (2017) High selfing
landraces from West-Africa: implication for conservation
rate inferred for white fonio [Digitaria exilis (Kippist.)
and breeding. Justus-Liebig University Giessen, Gießen
Stapf] reproductive system opens up opportunities for
Adoukonou-sagbadja H, Dansi A, Vodouhè R, Akpagana K
breeding programs. Genet Resour Crop Evol. doi:10.1007/
(2004) Collecting fonio (Digitaria exilis (Kipp.) Stapf, D.
s10722-017-0515-3
iburua Stapf) landraces in Togo. Plant Genet Resour Newsl
Blench RM (2012) Vernacular names for African millets and
139:63–67
other minor cereals and their significance for agricultural
Adoukonou-Sagbadja H, Dansi A, Vodouhe R, Akpagana K
history. Archaeol Anthropol Sci. doi:10.1007/s12520-012-
(2006) Indigenous knowledge and traditional conservation
0104-5
of fonio millet (Digitaria exilis, Digitaria iburua) in Togo.
Bouis HE, Welch RM (2010) Biofortification—a sustainable
Biodivers Conserv 15:2379–2395. doi:10.1007/s10531-
agricultural strategy for reducing micronutrient malnutri-
004-2938-3
tion in the global south. Crop Sci 50:S-20–S-32. doi:10.
Adoukonou-Sagbadja H, Wagner C, Dansi A et al (2007)
2135/cropsci2009.09.0531
Genetic diversity and population differentiation of tradi-
Brink M (2006a) Digitaria iburua Stapf. http://uses.plantnet-
tional fonio millet (Digitaria spp.) landraces from different
project.org/en/Digitaria_exilis_(PROTA)
123
Genet Resour Crop Evol
BrinkM (2006b) Secale cereale L. In: PROTA (Plant Resources review. Part 1. Problems. Int J Pest Manag 42:253–265.
of Tropical Africa/Ressources végétales l’Afrique doi:10.1080/09670879609372003
tropicale) Hilu KW, M’Ribu K, Liang H, Mandelbaum C (1997) Fonio
Brocklehurst PA (1977) Factors controlling grain weight in millets: ethnobotany, genetic diversity and evolution. S Afr
wheat. Nature 266:348–349. doi:10.1038/266348a0 J Bot 63:185–190. doi:10.1016/S0254-6299(15)30742-0
Bush SM, Krysan PJ (2010) iTILLING: a personalized approach Hong Y, Chen L, Du LP et al (2014) Transcript suppression of
to the identification of induced mutations in Arabidopsis. TaGW2 increased grain width and weight in bread wheat.
Plant Physiol 154:25–35. doi:10.1104/pp.110.159897 Funct Integr Genom 14:341–349. doi:10.1007/s10142-
Clément J, Leblanc JM (1984) Collecte IBPGR-ORSTOM de 014-0380-5
1977 au Togo. In: Prospection des Digitaria exilis (Fonio) Jansky S (2006) Overcoming hybridization barriers in potato.
en Afrique de l’ Ouest. ORSTOM, Marseille, France, Plant Breed 125:1–12. doi:10.1111/j.1439-0523.2006.
pp 3–7 01178.x
Clottey VA, Avornyo F, Addo-Kwafo A, Agyare WA (2006) Jideani IA, Jideani VA (2011) Developments on the cereal
The potential of fonio (Digitaria exilis Stapf) as feed for grains Digitaria exilis (acha) and Digitaria iburua (iburu).
monogastrics. Livest Res Rural Dev 18. http://www.lrrd. J Food Sci Technol 48:251–259. doi:10.1007/s13197-010-
org/lrrd18/7/clot18095.htm. Accessed 21 Feb 2017 0208-9
Dachi SN, Gana AS (2008) Adaptability and yield evaluation of Kahane R, Hodgkin T, Jaenicke H et al (2013) Agrobiodiversity
some Acha (Digitaria exilis and Digitaria iburua (Kippist) for food security, health and income. Agron Sustain Dev
Stapf) accessions at Kusogi-Bida, Niger State, Nigeria. Afr 33:671–693. doi:10.1007/s13593-013-0147-8
J Gen Agric 4:73–77 Kaoneka SR, Saxena RK, Silim SN et al (2016) Pigeonpea
Daniell H, Lin C-S, Yu M, Chang W-J (2016) Chloroplast breeding in eastern and southern Africa. Plant Breed
genomes: diversity, evolution, and applications in genetic 135:148–154. doi:10.1111/pbr.12340
engineering. Genome Biol 17:134. doi:10.1186/s13059- Kato Y, Yoshioka H, Iba R et al (2006) A newly bred rice variety
016-1004-2 ‘‘Minamiyutaka’’ for whole-crop silage. Bull Miyazaki
Dansi A, Adoukonou-sagbadja H, Vodouhe R (2010) Diversity, Agric Exp Stn 41:51–60 (in Japanese with English
conservation and related wild species of Fonio millet summary)
(Digitaria spp.) in the northwest of Benin. Genet Resour Kennedy G, Nantel G, Shetty P (2003) The scourge of hidden
Crop Evol 57:827–839. doi:10.1007/s10722-009-9522-3 hunger: global dimensions of micronutrient deficiencies.
Egan AN, Schlueter J, Spooner DM (2012) Applications of next- Food Nutr Agric 32:8–16
generation sequencing in plant biology. Am J Bot Khan S, Tariq R, Yuanlai C, Blackwell J (2006) Can irrigation
99:175–185. doi:10.3732/ajb.1200020 be sustainable? Agric Water Manag 80:87–99. doi:10.
Elshire RJ, Glaubitz JC, Sun Q et al (2011) A robust, simple 1016/j.agwat.2005.07.006
genotyping-by-sequencing (GBS) approach for high Konishi S, Izawa T, Lin SY et al (2006) An SNP caused loss of
diversity species. PLoS ONE 6:1–10. doi:10.1371/journal. seed shattering during rice domestication. Science
pone.0019379 312:1392–1396
FAO (2016) The state of food and agriculture. Rome Koreissi-Dembélé Y, Fanou-fogny N, Hulshof PJM, Brouwer
FAOSTAT (2017) Agricultural production, crop primary data- ID (2013) Fonio (Digitaria exilis) landraces in Mali:
base. In: Food and Agriculture Organization of the United nutrient and phytate content, genetic diversity and effect of
Nations. http://faostat3.fao.org/browse/Q/QC/E. Accessed processing. J Food Compos Anal 29:134–143. doi:10.
10 Feb 2017 1016/j.jfca.2012.07.010
Fekih R, Takagi H, Tamiru M et al (2013) MutMap?: genetic Koroch AR, Juliani HR, Simon JE (2013) Nutritional value of
mapping and mutant identification without crossing in rice. fonio (Digitaria exilis) from Senegal. In: African natural
PLoS ONE 8(7):e68529 plant products Volume II: discoveries and challenges in
Gigou J, Stilmant D, Diallo TA et al (2009) Fonio millet (Dig- chemistry, health, and nutrition. ACS symposium series,
itaria exilis) response to N, P and K fertilizers under pp 127–133
varying climatic conditions in west Africa. Exp Agric Krishna H, Alizadeh M, Singh D et al (2016) Somaclonal
45:401–415. doi:10.1017/S0014479709990421 variations and their applications in horticultural crops
Glaszmann JC, Kilian B, Upadhyaya HD, Varshney RK (2010) improvement. 3 Biotechn 6:54. doi:10.1007/s13205-016-
Accessing genetic diversity for crop improvement. Curr 0389-7
Opin Plant Biol 13:167–173. doi:10.1016/j.pbi.2010.01. Kuta DD, Kwon-ndung E, Dachi S et al (2003) Potential role of
004 biotechnology tools for genetic improvement of ‘‘lost crops
Godfray HCJ, Beddington JR, Crute IR et al (2012) The chal- of Africa’’: the case of fonio (Digitaria exilis andDigitaria
lenge of food security. Science 327:812. doi:10.4337/ iburua). Afr J Biotechnol 2:580–585
9780857939388 Larkin PJ, Scowcroft WR (1981) Somaclonal variation—a
He J, Zhao X, Laroche A et al (2014) Genotyping-by-se- novel source of variability from cell cultures for plant
quencing (GBS), an ultimate marker-assisted selection improvement. Theor Appl Genet 60:197–214. doi:10.1007/
(MAS) tool to accelerate plant breeding. Front Plant Sci BF02342540
5:484. doi:10.3389/fpls.2014.00484 Larson G, Piperno DR, Allaby RG et al (2014) Current per-
Hillocks RJ, Logan JWM, Riches CR et al (1996) Soil pests in spectives and the future of domestication studies. Proc Natl
traditional farming systems in Sub-Saharan Africa—a Acad Sci 111:6139–6146. doi:10.1073/pnas.1323964111
123
Genet Resour Crop Evol
Letunic I, Bork P (2016) Interactive Tree Of Life (iTOL): an friable callus of ‘‘Fonio’’ (Digitaria exilis (L.) Stapf). Sci
online tool for phylogenetic tree display and annotation. Hortic (Amst.) 125:494–499. doi:10.1016/j.scienta.2010.
Nucleic Acids Res 23:127–128. doi:10.1093/ 04.017
bioinformatics/btl529 Okuno A, Hirano K, Asano K et al (2014) New approach to
Li Q, Li L, Yang X et al (2010) Relationship, evolutionary fate increasing rice lodging resistance and biomass yield
and function of two maize co-orthologs of rice GW2 through the use of high gibberellin producing varieties.
associated with kernel size and weight. BMC Plant Biol PLoS ONE. doi:10.1371/journal.pone.0086870
10:143. doi:10.1186/1471-2229-10-143 Olodo KF (2016) Développement de nouveaux marqueurs
Li Y, Fan C, Xing Y et al (2011) Natural variation in GS5 plays microsatellites à partir du transcriptome du fonio chez les
an important role in regulating grain size and yield in rice. espèces cultivés (Digitaria exilis, Digitaria iburua) et
Nat Genet 43:1266–1269. doi:10.1038/ng.977 sauvage (Digitaria longiflora). Master thesis, Université
Lin Z, Li X, Shannon LM et al (2012) Parallel domestication of Cheikh Anta Diop de Dakar
the Shattering1 genes in cereals. Nat Genet. doi:10.1038/ Padulosi S, Thompson J, Rudebjer P (2013) Fighting poverty,
ng.2281 hunger and malnutrition with neglected and underutilized
Louwaars NP, de Boef WS (2012) Integrated seed sector species (NUS): needs, challenges and the way forward.
development in Africa: a conceptual framework for cre- Bioversity International, Rome
ating coherence between practices, programs, and policies. Patterson SE, Bolivar-Medina JL, Falbel TG et al (2016) Are we
J Crop Improv 26:39–59. doi:10.1080/15427528.2011. on the right track: can our understanding of abscission in
611277 model systems promote or derail making improvements in
Ma L, Li T, Hao C et al (2016) TaGS5-3A, a grain size gene less studied crops? Front Plant Sci 6:1268. doi:10.3389/
selected during wheat improvement for larger kernel and fpls.2015.01268
yield. Plant Biotechnol J 14:1269–1280. doi:10.1111/pbi. Peng S, Cassman KG, Virmani SS et al (1999) Yield potential
12492 trends of tropical rice since the release of IR8 and the
Mardis ER (2008) The impact of next-generation sequencing challenge of increasing rice yield potential. Crop Sci
technology on genetics. Trends Genet 24:133–141. doi:10. 39:1552–1559. doi:10.2135/cropsci1999.3961552x
1016/j.tig.2007.12.007 Ronald P (2011) Plant genetics, sustainable agriculture and
Mariac C, Scarcelli N, Pouzadou J et al (2014) Cost-effective global food security. Genetics 188:11–20. doi:10.1534/
enrichment hybridization capture of chloroplast genomes genetics.111.128553
at deep multiplexing levels for population genetics and Saha D, Gowda MVC, Arya L et al (2016) Genetic and genomic
phylogeography studies. Mol Ecol Resour 14:1103–1113. resources of small millets. CRC Crit Rev Plant Sci
doi:10.1111/1755-0998.12258 35:56–79. doi:10.1080/07352689.2016.1147907
Marouzé C, Fliedel G, Thaunay P, Cruz JF (2008) Designing a Saidou SIDI, Bakasso Y, Inoussa MM et al (2014) Diversité
fonio mill; screening an operating principle and its vali- agro-morphologique des accessions de fonio [Digitaria
dation. AMA Agric Mech Asia Africa Lat Am 39:9–15 exilis (Kippist.) Stapf] au Niger. Int J Biol Chem Sci
Massawe F, Mayes S, Cheng A (2016) Crop diversity: an 8:1710–1729
unexploited treasure trove for food security. Trends Plant Sanogo MD, Hash CT (2014) Enhancing fonio potential by
Sci 21:365–368. doi:10.1016/j.tplants.2016.02.006 intensifying production systems in partnership with farm-
McCallum CM, Comai L, Greene EA, Henikoff S (2000) Tar- ers of Mandela, Siguila and Kobougou. http://www.ccrp.
geted screening for induced mutations. Nat Biotechnol org/projects/fonio-i-ii. Accessed 08 Feb 2017
18:455–457. doi:10.1038/74542 Sarah G, Homa F, Pointet S et al (2017) A large set of 26 new
Meuwissen THE, Hayes BJ, Goddard ME (2001) Prediction of reference transcriptomes dedicated to comparative popu-
total genetic value using genome-wide dense marker maps. lation genomics in crops and wild relatives. Mol Ecol
Genetics 157:1819–1829 Resour 17:565–580. doi:10.1111/1755-0998.12587
Michael SJ, Davis F, Csuti B et al (1993) Gap analysis: a geo- Sasaki A, Ashikari M, Ueguchi-Tanaka M et al (2002) A mutant
graphic approach to protection of biological diversity. gibberellin-synthesis gene in rice. Nature 291:1–2. doi:10.
Wildl Monogr 123:3–41 1038/416701a
Morales-payán JP, Ortiz JR, Cicero J, Taveras F (2002) Digi- Scarcelli N, Barnaud A, Eiserhardt W et al (2011) A set of 100
taria exilis as a crop in the Dominican Republic. In: Janick chloroplast DNA primer pairs to study population genetics
J, Whipkey A (eds) Trends in new crops and new uses. and phylogeny in monocotyledons. PLoS ONE 6:e19954.
ASHS Press, Alexandria, pp 1–3 doi:10.1371/journal.pone.0019954
National Research Council (1996) Lost crops of Africa: volume Small E (2015) Teff & fonio—Africa’s sustainable cereals.
I: grains. The National Academies Press, Washington Biodiversity 16:37–41. doi:10.1080/14888386.2014.
Nelson GC, Rosegrant MW, Palazzo A et al (2010) Food 997290
security and climate change: challenges to 2050 and Song XJ, Huang W, Shi M et al (2007) A QTL for rice grain
beyond. International Food Policy Research Institute, width and weight encodes a previously unknown RING-
Washington type E3 ubiquitin ligase. Nat Genet 39:623–630. doi:10.
Ngozi UF (2013) The role of biofortification in the reduction of 1038/ng2014
micronutrient food insecurity in developing countries. Afr Stamp P, Messmer R, Walter A (2012) Competitive underuti-
J Biotechnol 12:5559–5566. doi:10.5897/AJB12.560 lized crops will depend on the state funding of breeding
Ntui VO, Azadi P, Supaporn H, Mii M (2010) Scientia Horti- programmes: an opinion on the example of Europe. Plant
culturae Plant regeneration from stem segment-derived
123
Genet Resour Crop Evol
Breed 131:461–464. doi:10.1111/j.1439-0523.2012. Varshney RK, Nayak SN, May GD, Jackson SA (2009) Next-
01990.x generation sequencing technologies and their implications
Su Z, Hao C, Wang L et al (2011) Identification and develop- for crop genetics and breeding. Trends Biotechnol
ment of a functional marker of TaGW2 associated with 27:522–530. doi:10.1016/j.tibtech.2009.05.006
grain weight in bread wheat (Triticum aestivum L.). Theor Vodouhè SR, Achigan Dako EG (2006) Digitaria exilis (Kip-
Appl Genet 122:211–223. doi:10.1007/s00122-010-1437-z pist) Stapf. In: PROTA (Plant Resources of Tropical
Takezawa S, Cisse M (2004) Domestication des Céréales au Africa/Ressources végétales l’Afrique tropicale). http://
Mema, Mali. In: Sanogo K, Togola T (eds) XIth Congress uses.plantnet-project.org/en/Digitaria_exilis_(PROTA).
of Panafrican Association prehistory and related fields, Accessed 6 Jan 2017
Bamako, 07–12 Feb 2001. Soro Print Color, Bamako, Vodouhe SR, Zannou A, Achigan Dako E (eds) (1998) Actes du
pp 105–121 premier atelier sur la diversité génétique du fonio (Digitaria
Taylor JRN, Schober TJ, Bean SR (2006) Novel food and non- exilis Stapf) en Afrique de l’Ouest, Conakry, Guinée, 4-6
food uses for sorghum and millets. J Cereal Sci août 1998. https://www.bioversityinternational.org/index.
44:252–271. doi:10.1016/j.jcs.2006.06.009 php?id=244&tx_news_pi1%5Bnews%5D=739&cHash=d3
Tilman D, Balzer C, Hill J, Befort B (2011) Global food demand 915d5b474597b6c3f120b2aa9bae18. Accessed 15 Feb 2017
and the sustainable intensification of agriculture. Proc Natl Vodouhè RS, Dako GEA, Dansi A (2007) Fonio: a treasure for
Acad Sci 108:20260–20264. doi:10.1073/pnas.111643 West Africa. Plant genetic resources and food security in
7108 West and Central Africa. Ibadan, Nigeria, pp 219–222
Vall E, Andrieu N, Beavogui F, Sogodogo D (2011) Les cultures Wu W, Liu X, Wang M et al (2017) A single-nucleotide poly-
de soudure comme stratégie de lutte contre l’insécurité morphism causes smaller grain size and loss of seed shat-
alimentaire saisonnière en Afrique de l’Ouest: le cas du tering during African rice domestication. Nat Plants.
fonio (Digitaria exilis Stapf). Cah Agric 20:294–300 doi:10.1038/nplants.2017.64
123
View publication stats