CHARACTERIZING FONIO PRODUCTION ACROSS SCALES:
INTERDISCIPLINARY INSIGHTS FROM A LOCAL FIELD TRIAL AND A SCOPING REVIEW
By
Abigail Dingus
A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
Crop and Soil Sciences- Master of Science
2022
ABSTRACT
As the changing climate places new stressors on agricultural production, novel and
adaptive management practices that provide resistance or resilience are needed to overcome
these challenges. Increased biodiversity can improve resilience and ecosystem function,
allowing agroecosystems to continue production after temporal or extreme climate events. One
way to increase both local and regional diversity is by broadening the number of crops
produced. Minor crops which suffer from lack of formalized research and are grown for cultural
value or local adaptations, may fill niches in agroecosystems outside their traditional production
systems. One of these minor crops is a small grain from West Africa called fonio. Fonio is
extremely valuable in West Africa for food security, nutrition, and cultural practices, but its value
has not yet been recognized globally. Increasing our understanding of this crop may provide a
pathway for improving fonio for its native habitat and new environments. While there are ample
uses of fonio and opportunities to improve production, there is limited research on fonio in most
disciplines. This research sought to fill knowledge gaps and inform future research on fonio
through two studies: a scoping review and a field study. A scoping literature review was
completed where research across disciplines was summarized and analyzed for gaps in
knowledge. The review found novel uses for fonio including building materials, pharmaceutical
uses, industrial uses, and feed for animals. Further use of fonio is limited by knowledge gaps in
the optimization of management for different environments, mechanization for production and
processing, and a lack of breeding efforts. To better understand the production and uses of
fonio more documentation is needed in field management, economics, seed systems, and
producer/consumer preferences. A major gap identified was the viability of fonio outside its
native habitat and the effect of fertilizer and seeding rate on fonio growth and forage quality. A
two-year (2021-22) field study conducted in Western Michigan tested fonio production in a new
environment, assessed forage quality, and determined the effect of seeding and fertilizer rate on
biomass and grain yields. Results indicated that low planting densities (4 kg/ha) improved grain
yields but reduced biomass. Fertilization greater than 19.5 kg N/ha increased biomass but had
no effect on grain yield. Grown in Michigan, fonio forage quality was high (RFQ=131-150) when
cut at booting stage but biomass yield was low compared to other summer annuals. From these
studies, we determined that fonio may be a useful crop in the US as a forage or cover crop,
filling a niche in perennial pastures during dry summer months. More research is needed to
further understand fonio management and global uses while ensuring West African producers
maintain sovereignty as the crop stewards of fonio.
ACKNOWLEDGEMENTS
These projects would not have been possible without the support of numerous advisors,
cheerleaders, and challengers. My advisors, Krista Isaacs, Kim Chung, Sieg Snapp, and Eric
Olson, were amazing sources of knowledge, guidance, and correction through the strenuous
process of starting and completing a thesis during a global pandemic. Krista Isaacs has gone
beyond her responsibilities as my advisor not only in these projects but also in my development
as a writer, a scientific thinker, and a lifelong learner. I would also like to acknowledge Jacob
Quincey at Fat Blossom farm for the invaluable knowledge he shared as well as the use of land,
equipment, and time devoted to the project. The Olson lab devoted resources such as a growth
chamber, greenhouse space, and training that greatly contributed to this project. I would like to
thank the members of the Snapp Lab, Cassida Lab, and the many friends who helped me plant,
harvest, thresh, collect data, analyze data, and edit this manuscript. Specifically, Joe Paling was
instrumental in training and advising me on the forage quality work and Ann Bybee-Finley on the
statistical analyses. This work was made possible by funding from the Plant Science Fellowship
from Michigan State University and generous donations from The Carter Harrison Endowed
Fund.
iv
TABLE OF CONTENTS
1. INTRODUCTION……………………………………………………….………...…………...……...1
2. RESEARCH QUESTIONS………………………………………………..……………....…..……..7
3. OPPORTUNITIES TO IMPROVE FONIO PRODUCTION: A SCOPING REVIEW TO
INFORM FUTURE RESEARCH…………………..…………………..……………8
4. GROWING FONIO IN MICHIGAN: THE EFFECT OF SEEDING AND FERTILIZER
RATES ON FONIO GRAIN YIELD, BIOMASS PRODUCTION, AND
FORAGE QUALITY…………………………………………….……..…………….54
5. CONCLUSIONS………………………………………....………….…………………....………….77
REFERENCES………………………………………...…….………..…….……….………………….79
v
1. INTRODUCTION
The changing climate is increasing biotic and abiotic stress on agroecosystems. Abiotic
stressors associated with climate change such as increased temperatures, temporal drought,
and reduced soil carbon limit overall moisture available to plants, impacting growth and yield of
crops (Aydinalp & Cresser, 2008). Increased strength of weather events is eroding soils of
essential nutrients, making continuous production more difficult. More frequent flood events and
rising sea levels are also damaging crops and permanently reducing usable cropland (Aydinalp
& Cresser, 2008). Abiotic stressors such as increased CO2, temperatures, and reduced soil
fertility may also increase pressure from weeds and other pests (R. M. Adams et al., 1998). As
stressors increase, agriculture production will likely decrease, causing economic loss and
negatively impacting food security.
It is essential for agricultural systems to adapt or quickly recover from extreme climatic
events. The ability for ecosystems to maintain organizational structure and production after a
disturbance is called resiliency (Holling, 1973) and is increased by management practices such
as crop diversification, use of local diversity, soil fertility management, and water conservation
(Altieri et al., 2015). Increased biodiversity is an essential way to build resilient agroecosystems
(Lin, 2011; Vandermeer et al., 1998) by improving ecosystem function. As functionally diverse
species use different niches in the soil, available resources are more fully utilized (Lin, 2011;
Song et al., 2014). Long term studies have shown that increasing plant biodiversity, specifically
with functional diverse groups such as forbes, legumes, and grasses helps build soil nutrients
and overall soil fertility compared to monocultures (Furey & Tilman, 2021). Efficient resource
use improves ecosystem function and builds soil fertility, creating long-term resilient
agroecosystems.
Closely related to resiliency is the concept of redundancy. Redundancy or insurance is
when one crop fails, and others may survive to produce a crop. Increasing the number of
species grown in one system can achieve this by providing a buffer from climatic variation (Lin,
2011). For example, in a warm-season intercrop study, mixtures of four species exhibited
greater yield stability across environments compared to monocultures of the same species
(Bybee-Finley et al., 2016). Increased number of species by crop rotations, intercropping, or
integration of livestock can buffer against stressors from climate change and complete
economic loss for farmers.
As abiotic stressors increase, crops are vulnerable to effects of disease and pests.
Increased biodiversity may also help decrease biotic pressure from climate change. Specifically,
1
using crops or varieties which are resistant to the pest or disease can reduce the spread and
pressure (Lin, 2011; Reiss & Drinkwater, 2018). In Florida, fall army worm infestations were
reduced 70-96% by changing the cover crop from a monocrop of army worm susceptible
sorghum sudangrass to a mixture of sun hemp and legume cover crop (Meagher et al., 2022).
In China, the fungal infection known as rice blast was reduced on the field scale and yields were
maintained by intercropping varieties resistant to rice blast and lower yields with susceptible,
high yielding varieties (Revilla-Molina, n.d.).
The management practice of increasing biodiversity builds resiliency, increases
redundancy, and creates resistance to biotic stressors and is implemented across both temporal
and spatial scales. On a temporal scale, increased number of rotations can improve yield
stability. In Nebraska, diverse rotations of 2-4 years improved yield stability of corn and sorghum
yields compared to less diverse rotations (Sindelar et al., 2016). The variety of crops grown in
monoculture through a single year by using cover crops may also increase resilience and yields
by improving soil fertility (Thapa et al., 2021). Forms of intercropping such as relay cropping,
and agroforestry increase biodiversity both spatially and temporally. Spatial diversity may be
improved on the field, farm, or regional scale. In the field, intercropping different species
together can build resiliency and yield stability across agroecosystems (Bybee-Finley et al.,
2016). At the farm level, diversity may be increased by incorporating more variety of crops,
integrating livestock and pastures onto the farm, or by developing riparian zone or buffer zones.
Regional diversity may also be increased using non-native crop species which are non-invasive.
The use of minor crops may be a way to protect global diversity and increase local and
regional biodiversity. Minor crops are plants that are not domesticated or semi-domesticated
and are used for medicines, nutrition, fuel, and other cultural practices. They are called “minor”
or “neglected crops” because they have undergone “incomplete selection” or had limited formal
breeding, leading to challenges such as shattering, small grain size, or lodging; or they may
have other hurdles, such as processing, that require extra labor. They are often locally adapted,
resistant to stressors, and maintain an abundance of new genetic material (Dansi et al., 2012;
Padulosi et al., 2011). However, they are rarely utilized beyond their local context. They also are
often replaced by new, higher yielding varieties of major crops, putting them at risk of genetic
loss or extinction.
The use of minor crop species has benefitted U.S. agricultural systems by increasing
ecosystem function, redundancy, and reducing pest pressure. For example, crabgrass (Digitaria
sanguilis), typically considered a weed in the United States, is being grown more widely as a
forage crop. The use of forage crabgrass was shown to increase the resiliency of U.S. pastures
2
during times of drought due to increased water use efficiency compared to other perennial
forages (Gelley et al., 2020). The use of the minor crops, cowpea and sun hemp as cover crops
in southern Texas and Florida were shown to reduce the pressure of fall army worms (Meagher
et al., 2022). Cover crop mixtures including minor crops like cowpea, sun hemp, pearl millet,
and sorghum sudangrass increased yield stability across environments (Bybee-Finley et al.,
2016). These studies have shown that the use of minor crops provided more biodiversity and
increased resiliency in agroecosystems. A minor crop which has not yet been studied in the
United States is fonio (Digitaria exilis), a resilient and diverse small grain crop from West Africa
which may greatly contribute to the resiliency of Michigan agroecosystems as a cover, forage,
or grain crop.
Fonio (Digitaria spp.) is a minor grain crop with essential uses throughout West Africa
also commonly called “acha” or “fundi”. In many regions, it is used for food security as quick
maturing varieties fill the “hunger gap” when food stores from the previous year are low and
before the major harvest (Fogny-Fanou et al., 2010a; Vall et al., 2011b). Fonio is highly adapted
to low fertility soils and has high tolerance for drought. Often, it is produced at the end of many
crop rotations, when the soil is exhausted (Kanlindogbe et al., 2020). Fonio is known to have a
negative effect on the lifecycle of certain infectious fungus species such as charcoal-rot
(Macrophomina phaseolina) (Ndiaye & Termorshuizen, 2008) and nematodes like root-knot
worms (Meloidogyne javanica) (Sarr & Prot, 1985) making fonio advantageous not only for food
security, but also for integrated pest management in Africa (Ndiaye & Termorshuizen, 2008).
Fonio yields range between 200-1200 kg/ha and have generally increased over the last 50
years (Figure 1.1), but in most countries, total production area remains low (below 80,000 ha).
The exception to this is Guinea which has higher yields and increased production area greatly in
the last 50 years for unknown reasons. Depending on the region, fonio is eaten steamed like
couscous and accompanied with a sauce, as a snack, as porridge for breakfast, or as baked
goods, and it has the potential to be used in a large array of items such as breads or crackers (I.
A. Jideani, 1999). The high protein and fiber content make it a nutritious grain for humans and
livestock (C. I. Ukim et al., 2021). Fonio also has significant cultural value and is prepared for
special occasions, celebrations, and ancestral worship (Adoukonou-Sagbadja et al., 2008;
Ayenan et al., 2018).
3
1200
1000
Average Yield (kg/ha)
800
600
400 Benin
Burkina Faso
200 Guinea
Guinea-Bissau
Mali
0 Niger
1970 1980 1990 2000 2010 2020
Nigeria
600000
500000
Production Area (ha)
400000
300000
200000
100000
0
1970 1980 1990 2000 2010 2020
Decade
Figure 1.1: Yield and production areas, averaged across decade, of seven major fonio
producing countries in West Africa. Data Source: FAO Stat
Fonio is one of the oldest indigenous grains in West Africa, but it is not widely known by
the rest of the world and may have potential to impact several disciplines (Cruz et al., 2016).
Fonio’s adaptability and genetic diversity has potential to contribute to global food and nutritional
security in a changing climate. Advantageous traits like drought resistance and low nutrient
requirements are attributed to an extensive root system and reliance on mycorrhizal networks
(Ndoye et al., 2016). Fonio may also be bred for a variety of different climates and growing
conditions because of its preserved genetic diversity. For example, in Togo alone, 42 different
4
varieties were characterized and collected from traditional farmers with varying attributes
(Adoukonou-Sagbadja et al., 2008). Fonio is also considered a health food that has a low
glycemic index and in gluten content while being high in essential amino acids such as lysine
and methionine (Cruz et al., 2016). Thus, fonio may be advantageous for the rest of the world
as a climate smart crop, as a genetic resource, or as a highly nutritious food.
Despite the many advantages to growing this crop, there are still many barriers to its
production. Fonio can be variable in yield (200-1200 kg/ha) due to small grain size and grain
loss (Cruz et al., 2016). Grains can be lost due to shattering (up to 30%), lodging, and
postharvest processing (Ayenan et al., 2018; Cruz et al., 2016). The common weed, striga, can
also cause significant yield loss (Ayenan et al., 2018; Kanlindogbe et al., 2020). Over time these
barriers may continue to decrease fonio production as it is replaced by crops such as sorghum
or maize that are higher yielding and require less labor (Dansi, Adoukonou-Sagbadja, &
Vodouhe, 2010). In Senegal, farmers have recognized the decline in fonio production mostly
due to lack of seed, tedious production practices, and disinterest of young farmers (Diop et al.,
2018). The decrease in production is thus a complex social, cultural, and technological issue.
With the decreased production of fonio, it may continue to be neglected by modern science
unless steps are taken to preserve this valuable crop.
Because of the lack of research on fonio, there are gaps in the literature that still need to
be filled and many disciplines can contribute to the optimized production of fonio. For example,
engineers may be consulted for the mechanization of harvest and threshing of fonio. Studying
the genetics and trait heritability may substantially improve fonio as a crop and management
practices may also overcome barriers such as low yield and yield loss. For example, optimizing
fertilization practices has potential to decrease pest pressure such as striga (Gigou et al., 2009;
Kanlindogbe et al., 2020). It has also been found that low levels of fertilization (30 kg/ha) can
increase yields by 22% and increase grain protein content (Gigou et al., 2009). With investment
in crop and production methods, significant increase in yield may be achieved and costs
reduced. Disciplines such as nutrition, animal science, and material science may also contribute
to the development of this crop by proposing novel and profitable uses. Economists may identify
optimal investment strategies to improve profitability, potentially driving research and continued
production by West African farmers. Many of these disciplines have contributed to our
knowledge of fonio biology, production, and uses, but there has been a lack of integration
between disciplines.
Testing fonio in new environments and cropping systems such as in the United States
may spread awareness about the climate smart advantages, potentially creating new markets
5
and driving innovation. However, to our knowledge, fonio has never been grown in the United
States. There is some research on production methods in West Africa, but there has never been
research on how these methods may translate to new climates and cropping systems. One
location that may benefit from the drought tolerance of fonio is in Michigan cropping systems.
Specifically in western Michigan, the climate and the soil conditions are different from many
places fonio is grown in West Africa. Michigan averages 15 ˚C night time lows, 25.5 ˚C daytime
highs, and 306 mm precipitation during the growing season often with temporal drought (June-
September) (Temperature, Rainfall and Degree Day Summary - MSU - Enviroweather, n.d.).
Because of unpredictable and worsening summer drought, fonio, being a drought tolerant crop,
may contribute to several Michigan agricultural systems and increase overall spatial diversity in
Michigan. To test these potential niches for fonio, it must be tested in Michigan growing
conditions and production practices need testing like seeding rates and fertilization strategy.
Climate change is challenging agricultural production across the globe. Increasing
biodiversity and utilizing minor crops will improve the ability of agroecosystems to overcome
stressors. Fonio, as a minor crop, may contribute to biodiversity in cropping systems but has
been under-researched and may continue to be neglected as farmers replace fonio with major
crops. Many disciplines have contributed to our knowledge of fonio but there has been lack of
integration between disciplines. Thus, there are insights to be gained from mapping all research
on fonio and integrating those findings across disciplines. Testing fonio in new environments
and new cropping systems may also spread awareness about the climate smart advantages,
but fonio production must be studied in new environments. Specifically, basic management
practices such as fertilization and seeding rates are needed to start growing fonio globally. The
purpose of this research is to characterize fonio production in a new environment, studying
specific production parameters and mapping the extent of research that has been completed on
fonio to prepare for future research.
6
2. RESEARCH QUESTIONS
Scoping Review Research Questions and Hypotheses
1. What are the existing knowledge and knowledge gaps in fonio literature in terms of its uses,
production, and breeding?
2. What are specific improvements needed to optimize fonio production? What are drivers for
innovation in fonio research?
We hypothesize that a scoping literature review integrated across disciplines like agronomy,
economics, and plant sciences will construct a clear, informed plan to improve fonio as a crop
and the technology needed to increase production. We predict that the major improvements
needed will be widespread mechanization of production, breeding improvements for increased
yields and processing qualities, and optimized production practices.
Field Study Research Questions and Hypotheses
1. How does fertilizer rate affect lodging, total biomass, grain yield, and plant growth
characteristics?
2. How does seeding rate affect lodging, total biomass, grain yield, plant growth characteristics,
and forage yield and quality?
We hypothesize that increased fertilizer will increase total biomass by increasing growth
parameters such as height and number of tillers but may increase lodging and not improve
overall grain yield due to plant partitioning. For seeding rate, we hypothesize that increased
seeding rate will increase total biomass and grain yield but may decrease growth parameters
such as plant height and tillers as competition increases. We hypothesize that seeding rate will
not affect forage quality but will increase forage yield.
7
3. OPPORTUNITIES TO IMPROVE FONIO PRODUCTION: A SCOPING REVIEW TO
INFORM FUTURE RESEARCH
3.1 Abstract
Use of minor crops can increase biodiversity, improving resiliency and stability of
agroecosystems as stressors from climate change increase. Fonio (Digitaria exilis) is a minor
grain crop in West Africa valued for its tolerance to extreme conditions like drought and poor
nutrient soils and for its nutritious grains which are gluten free and high in protein. However, little
formal research has been conducted on fonio. The objective of this scoping review was to
summarize all electronically published research on fonio using an agroecological lens and
focusing on biophysical, social, and economic elements to determine key knowledge gaps in
production to direct future research. The review found numerous novel food and non-food uses
for grain and waste products which may increase demand for fonio. Production of fonio is still
limited with numerous knowledge gaps, particularly in the relationship of fonio response to
management, genetics, and environment. Mechanization is also needed to reduce labor
requirements, specifically for women. The identification of markers and study of mutagenesis in
fonio lay the groundwork for breeding advancements which could increase production. Greater
documentation of fonio producers’ and consumers’ preferences is needed to inform breeding
objectives. Fonio research must increase to realize its full potential, but development of this crop
must acknowledge and protect the stewardship of the West African farmers who cultivate and
protect this valuable resource.
3.2 Introduction
Agricultural production is challenged by increased stress from climate change. Climatic
events like drought and storms are worsening, causing increased pressure from pests and
diseases. To overcome these challenges, agroecological management practices can help
systems resist stressors or maintain production during extreme climate events or with changing
temperatures. A well-researched management practice to increase resiliency, redundancy, and
stability in agroecosystems is increasing biodiversity (Altieri et al., 2015; Furey & Tilman, 2021;
Lin, 2011). A review of 172 cases revealed that increased crop diversity contributed to resilience
through improved ecosystem function and preservation, sustainable use of natural resources,
and utilization of stress-tolerant crops (Mijatović et al., 2013). Biodiversity may be improved on
multiple scales through practices such as cover cropping, intercropping, livestock integration, or
increased number of rotations and crop species (Bybee-Finley & Ryan, 2018; Sindelar et al.,
2016; Thapa et al., 2021). Broader use of minor crops which are non-native, non-invasive is one
way to increase the total number of crops species. Minor crops are often semi-domesticated
and used for medicines, nutrition, or cultural practices. While the widespread use of these crops
suffers from lack of formalized research or breeding efforts to address production and/or
8
processing challenges, they are still grown because of their cultural value or adaptations to local
stressors (Dansi et al., 2012; Padulosi et al., 2011).
A minor crop with little formalized research is fonio (Digitaria exilis), a small grain
cultivated in West Africa. Fonio increases crop diversity in West Africa since it is produced as
the last crop in rotation when the soil is exhausted (Kanlindogbe et al., 2020). As a drought
tolerant crop, fonio contributes to the resiliency of the food system as quick maturing varieties fill
the hunger gap when food stores from the previous year are low and before the major harvest
(Fogny-Fanou et al., 2010a; Vall et al., 2011b). It also provides diversity in animal feed when
leftover straw is recycled to feed cattle and other animals (Cruz et al., 2016). The crop is deeply
embedded in culture and food traditions as a food regularly prepared for special occasions,
celebrations, and ancestral worship (Adoukonou-Sagbadja et al., 2008; Ayenan et al., 2018).
Fonio is already benefitting West African farmers through traditional value, nutrients, and food
security.
Globally, these agroecological advantages have not yet been recognized, but fonio, as
a grain or forage crop, has enormous potential. The genetic diversity of fonio may also have
global benefits for food and nutritional security. Similarly, a nutritious pseudo-grain originating
from the Andes, quinoa, has an abundance of genetic diversity and is bred for different
agroecosystems, with potential to increase food security globally (Bazile, Pulvento, et al., 2016;
Jacobsen, 2003). In Togo alone, 42 varieties of fonio were characterized (Adoukonou-Sagbadja
et al., 2008). Filling specific niches with fonio may also increase resilience of agroecosystems.
For example, crabgrass (Digitaria sanguilis) is considered a weed in the United States but is
grown as a forage crop in southern US and increases the resiliency of US pastures during times
of drought due to its increased water use efficiency (Gelley et al., 2020). There is potential to
find both diverse and economic uses. For example, fonio may increase global crop diversity and
economic output as a second crop since there are very fast maturing varieties or as a dual use
crop for both grain and forage.
While there are ample uses of fonio and opportunities to improve production, there has
been limited research. The existing literature spans multiple disciplines but generally there is
very little research on fonio within or across disciplines. Interdisciplinary analysis and
engagement could lead to creative and viable solutions to challenges in fonio production. For
example, efficiencies may be improved by consulting engineers for the mechanization of harvest
and threshing of fonio, reducing labor demands on women, and increasing household capacity
to produce more fonio. With improved harvesting and processing methods, a significant
reduction in women’s labor and increase in yield may be achieved. Furthermore, co-creation of
9
knowledge by understanding the nuances of farmer trait preferences in combination with
studying the genetics and trait heritability may substantially improve fonio as a crop. Additional
insights may be found through interdisciplinary engagement to assess if genotype by
environment by management influences the nutrient quality of end-products for both humans
and livestock. Disciplines such as nutrition, animal science, and material science may also
contribute to the development of this crop by proposing novel and profitable uses. Many
disciplines contribute to our knowledge of fonio but there is a lack of integration between
disciplines. We propose to examine these gaps through a scoping review to gain insights from
the integration of research across disciplines.
Scoping reviews are often used to identify gaps in knowledge across a broad range of
disciplines (Arksey & O’Malley, 2005). These reviews are common in the health sciences where
a vast amount of research must be synthesized to find gaps in knowledge (Tricco et al., 2018).
In the case of fonio, there is very little primary research, and it is spread across many
disciplines. Several systematic reviews have been done within disciplines such as nutrition
(Kanupriya & Dhiman, 2021; Salahudeen & Orhevba, 2021), food science (Zhu, 2020), breeding
(Abdul & Jideani, 2019), and genetics (Ayenan et al., 2018). There are no known reviews for
agronomy, animal science, engineering, or plant physiology, or across disciplines. Here, we use
a scoping review to map the areas of fonio research which have been studied, highlight
knowledge gaps and opportunities related to fonio production, and analyze how production may
be impacted by multiple disciplines (Arksey & O’Malley, 2005; Tricco et al., 2018). The
objectives of this scoping review are to:
1. Build a database and summarize all published research on fonio
2. Identify knowledge gaps in fonio production
3. Analyze how multiple disciplines may inform production knowledge gaps
This research is needed to better inform breeders, engineers, and farmers. Learning
from other disciplines and crops may prevent harming the traditional farmers like in the case of
quinoa (Drew et al., 2017) or technology that is unlikely to be adopted like in the case of
sorghum breeding (Diallo et al., 2018). A cross-disciplinary literature review of fonio will increase
our knowledge about fonio and provide insights for future research.
10
3.3 Theoretic Framework
An agroecological lens was used to analyze key disciplines related to fonio production.
Agroecology has ten elements summarized by the FAO’s publication, The 10 Elements of
Agroecology (FAO, 2018), and include: diversity, co-creation of knowledge, synergies,
efficiency, recycling, resilience, human and social value, culture and food traditions, responsible
governance, and circular and solidarity economy. This theory integrates elements from multiple
disciplines, specifically agronomy, ecology, sociology, and economics and is widely accepted
framework for interdisciplinary analysis of agricultural systems (Dalgaard et al., 2003). Scoping
and reviewing literature on fonio production benefits from this framework by going beyond
biophysical systems to the social and economic aspects of crop research and breeding. Specific
aspects of the agroecological framework like soil health, animal health, biodiversity, social
value, and food traditions, were directly summarized as related to fonio in specific disciplines.
We also emphasize specific topics which would benefit from co-creation of knowledge such as
breeding objectives and development of an equitable seed system.
To guide our analysis from the generation of specific, basic knowledge and the
application of that knowledge, we simplified the ten elements into three broad categories. Three
overlapping areas were the biophysical, social, and economic aspects of each topic (Figure
3.1). We used these categories to evaluate what basic knowledge exists and how it may be
applied with potential effects on social and economic aspects. For example, there are proposed
uses for fonio and the physical attributes of fonio have been studied to allow for these uses, but
there are unknowns on the application such as: How do the fonio products compare
economically to the original product and how may the new uses for fonio affect the producers?
Also are the new products acceptable to consumers and how has adoption of fonio changed
over time? These proposed applications for fonio may then direct breeding programs and other
biophysical research.
Social
Knowledge Application
Biophysical Economic
Figure 3.1: Graphical depiction of the framework used to analyze each topic area for
research gaps moving from basic knowledge to application of knowledge.
11
3.4 Methods
A scoping review of literature on fonio was conducted across disciplines to map existing
literature, identify key knowledge gaps, and integrate those findings. The methodological
framework proposed by Arksey & O’Malley (2005) was used by including the five stages the
authors identify for conducting a scoping literature review: identify the research question,
identify relevant studies, study selection, chart the data, and summarize and report the results
(Figure 3.2). The 20 elements for a scoping review created by Tricco et al. (2018) were also
included. As with many scoping reviews, our purpose was to review and synthesize across
disciplines rather than evaluate the scientific quality of each study included (Arksey & O’Malley,
2005). A broad research question was identified: what is known from the existing literature
about fonio; its biology, production, and uses? The parameters of study were further defined by
narrowing the review to true fonio (digitaria exilis). A similarly related species called black fonio
(digitaria iburu), is often included and compared in research on true fonio. Iburu or black fonio
was included in the study, but not specifically searched for. Other crab grass species (Digitaria
spp.), and animal or wild fonio (Brachiaria deflexa) (Lewicki, 1974) were excluded from the
review. Black fonio or iburu is closely related and has similar uses to fonio but is rarely studied
exclusively. Digitaria exilis was chosen since it is the most widespread through West Africa
(Cruz et al., 2016).
To identify relevant studies, two primary search engines were used: CAB abstracts and
Google Scholar. The search terms “fonio”, “acha”, and “digitaria exilis” were used in each
database which yielded a total of 1,258 references, last searched on 1 June 2022. Search
parameters were not limited by date to track fonio research through time. All languages were
included since most fonio research from Africa is published in French. Articles were translated
by google translate or found in an English version. Conference reports and abstracts were
excluded because the information published in articles and conference reports/abstracts was
almost always duplicated. Reference lists from extensive reviews (Cruz et al., 2016) were also
searched. Only electronic published literature was included. It is possible that research not yet
electronically published may have been overlooked, but due to practical constraints authors did
not hand search key journals or contact relevant organizations as suggested by Arksey &
O’Malley (2005). No other restrictions were utilized, but sources were sorted by date for ease of
extraction.
12
Figure 3.2: Methodology for the scoping literature search and selection
All references were screened by title and abstract and added to a database. Unrelated,
duplicate, or conference papers were eliminated yielding a total of 320 references which were
added to a Zotero reference manager. Many of the irrelevant references were related to health
fields using the abbreviation “ACHA” for an organization or for the last name of an author.
Articles that referenced fonio but did not do research directly related to fonio were also
excluded. Next, the data were extracted and added to an excel datasheet (“charting the data”
according to Arksey & O’Malley, 2005). The data included for each reference were author(s),
year of publication, title, study country, discipline, subtopic, aims, methodology and important
results taken directly from the abstract. Discipline criteria were based on Table 3.1. Subtopics
13
were generated iteratively by reading abstracts in each of the disciplines and grouping similar
topics together.
Table 3.1: Description of the disciplines which the literature was sorted
Discipline Details
genetics of fonio including potential genetic tools for
progressing the breeding of fonio, the origins of digitaria,
Genetics
may contribute to conservation and understand diversity with
Biology implications for breeding
microscopic structure and function of fonio or its metabolic
Physiology reactions, enzymes, germination, or processes and growth
affecting moisture content
the actual process of growing and cultivating fonio, also
Agronomy referring to the crop systems or distribution of fonio
production
Breeding methods of breeding or information on varieties or traits
Production related to pests and disease that affect fonio or ways fonio is
Pathology
resistant or could produce a solution for pest and diseases
Economic analysis of fonio production or issues of value
Economics
chain
the optimization or creation of fonio mechanization for
Engineering
production and processing
the properties of fonio incorporated building materials or
Material
properties of starch from fonio and possible food,
Science
pharmaceutical, and industrial uses
Animal Science uses as feed or fodder for animals
Uses nutritional benefits, chemical composition of fonio, including
Nutrition
things mentioning the amino acid, protein, gluten content
consumption information, impact of fonio on foods, research
Food Science related to starch properties, flour and flour mixture
properties, or the fermenting properties of fonio grains
synthesis of information on fonio or other cereal crops, no
Reviews new study or discovery, a summary of current research or
findings
After the database was created, subtopics and disciplinary findings were summarized.
Each reference’s abstract was read and paraphrased. Then, all paraphrases associated with the
subtopic were read and a summary was written. From each subtopic in “Uses”, a summary table
of all potential products from fonio was created and summarized. From each subtopic in
“Production”, a list of major research findings and knowledge gaps were recorded. This process
included a full reading and extraction of key papers to understand production practices and
14
conditions more fully. Synergies between disciplines were then explored. For example, from
engineering, it may be found that threshers often crush fonio grains and thus it is difficult to
develop mechanized threshing that dehulls the grain without crushing it. This finding may
influence the discipline of breeding which could select for harder grains or more easily hulled
fonio grains. From each subtopic in “Biology” key findings related to fonio production or breeding
were added to the above section to summarize and find key gaps in our knowledge of fonio
physiology related to production.
3.5 Results
3.5.1 Research Overview
The first published record of fonio was in 1934 when it was reported as a “foodstuff” in
early dietetics research (Turner, 1934). Since then, research has steadily increased and
expanded to more disciplines (Figure 3.3). The topic area of “Uses” for fonio including the
disciplines of Nutrition, Food Science, Animal Science, and Materials Science are the most
studied areas. A total of 64% of the primary research is focused on the uses of fonio. Over time,
fonio research has increased especially in categories of uses and biophysical knowledge.
Research on production has continued but has not significantly grown compared to the other
categories.
15
160
Review
140
Material Science
120 Food Science
100 Nutrition
Animal Science
Count 80
Engineering
60 Economics
40 Pathology
Agronomy
20
Physiology
0 Breeding
1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s 2010s 2020s Genetics
Decade
30
30
25
25
20
20
Count
15 15
10 10
5 5
0 0
1930s 1940s 1950s 1960s 1970s 1980s 1990s
Decade Year
Figure 3.3: Distribution of the references through time by discipline. The disciplines are
grouped into three topic areas indicated on the figure by color pallet: Biophysical elements
(blue), Production (green), and Uses (Orange). The top graph counts the number of papers
compared to decade except for the 2020’s which only include data from two years. The bottom
figures expand the 1900s to see specific topic breakdowns (left) and expand the 2000s by year
to demonstrate the year by year growth of fonio research (right).
16
The earliest publications on fonio are found in descriptive studies of West African diets
or farming practices (Murdock, 1960; Porters, 1955; Turner, 1934). Fonio (Digitaria exilis) has
been included in many more reviews and descriptive studies since the mid-1900s mostly
included as a minor crop grown in West Africa (Supplemental Table 1). A notable exception
was a description of the history and cultivation of fonio or funde in the Dominican Republic,
summarized by Morales-payán et al. (2002) where fonio is eaten as an aphrodisiac. Another key
publication is: Fonio: An African Cereal, where Cruz et al. (2016) synthesized the distribution,
production practices, and mechanization of fonio. Very few reviews have summarized original
research about fonio (Abdul & Jideani, 2019; Ibrahim Bio Yerima & Achigan-Dako, 2021; I. A.
Jideani & Jideani, 2011; Kanlindogbe, Sekloka, Amégnikin Zinsou, et al., 2020) and to the
authors’ knowledge, there have been no reviews written to synthesize research across
disciplines.
3.5.2 Fonio Uses (Table 3.2)
Culinary Uses
With a nutty flavor and multiple culinary uses, fonio is eaten across different West
African cultures and has potential to replace major grains. Traditionally, fonio is eaten as fluffed
couscous with sauce or as nutritious porridge. These porridges are suitable as a weaning food
based on the nutrient content and sensory qualities (Eboagu et al., 2020; Ugwuona et al., 2012).
Foods traditionally made with maize or small grains replaced with fonio have generally high
acceptability. Added to soups, fonio increased overall nutrient value of traditional foods like
pumpkin soup (Helen & Ihekanu, 2009). Fonio flour can make gluten free baked goods with
functional baking properties improved with the addition of 4% carboxymethylcellulose (CMC) or
3% xanthum gum (C. e. Chinma et al., 2015; V. A. Jideani et al., 2007). Mixing fonio into other
flours may also increase nutrient content while preserving functional properties (Supplemental
table 2). As with other small grains, fonio may be used to make traditional fermented or malted
drinks but may not be economical for brewing (Nzelibe et al., 2000; Nzelibe & Nwasike, 1995).
Many of these products are considered acceptable based on nutrient content and sensory
characteristics like taste, texture, and smell ranked by panelists. Despite the increased research
on food uses for fonio, there is little information on changing fonio consumption and perception.
Research on fonio consumption is limited but there is evidence of differences between
rural and urban settings for fonio consumption. In capital cities of Mali, Burkino Faso, and
17
Guinea, consumer surveys representative of the population reported that the majority of people
enjoyed eating fonio and wished to increase consumption (Konkobo-Yameogo et al., 2004). In
another survey in urban Mali, two-thirds of women reported eating fonio monthly (Fogny-Fanou
et al., 2010b). In these urban areas, fonio is valued as a health food or as a food for festivals,
but consumption is limited by cost and a lack of easy-to-make fonio products like par-boiled or
fully processed (Fogny-Fanou et al., 2010a; Konkobo-Yameogo et al., 2004). In rural areas of
Mali, Vall et al. (2011) found through sales analysis and farmer surveys, that fonio is bought
more often during lean periods and is used for seasonal food security. They found that the
constraint to consumption was not cost but a lack of desire to consume fonio when other, more
modern grains are available. Similarly in rural Senegal, fonio farmers most often indicated that
fonio was important during food shortages (Diop et al., 2018). Unlike consumption in urban
areas, fonio may be consumed in rural settings more often out of caloric need. Because of
limited documentation on consumption and perception of fonio, it is unknown if these patterns
between rural and urban areas are widespread across West Africa and how the perception may
affect increased use or adoption of fonio products.
Table 3.2: Uses for fonio across disciplines
Category Product Details Citation
Drinks Kunu-Zaki A drink made in Nigeria traditionally (J. A. Ayo, 2004;
Drink made with millets but was acceptable Oluwajoba et al.,
to a panel in aroma, taste, and 2013)
nutritional qualities when made with
fonio.
Malted Drink A variety of drinks made in W. Africa (Abdulquadri &
from a variety of grains. Panels Rahman, 2017;
deemed a malted fonio drink Badejo et al.,
acceptable in taste and nutritients with 2017)
additives like tugernut extract.
Fermented Brewing fonio as a mixture increased (Nzelibe et al.,
drink the brew’s sensory qualities. Five 2000; Nzelibe &
varieties of fonio were identified to Nwasike, 1995)
have good brewing qualities.
18
Table 3.2 (cont’d)
Food Baby/Weaning Fonio porridge as a weaning food was (Eboagu et al.,
Food acceptable in nutrient quality, flavor, 2020; Ugwuona
and color. et al., 2012)
Porridge As a breakfast food or a porridge, (Henry-Unaeze,
fonio was high in fat, protein, calcium, 2011; Mbaeyi-
and was acceptable in taste. Nwaoha &
Uchendu, 2016)
Masa A fermented rice cake eaten in Nigeria (Malomo &
acceptable in nutrients and taste when Abiose, 2020)
rice was 20% substituted with fonio
and 20% substituted with soy.
Ogi and Agidi A fermented rice or corn pudding (Chisom et al.,
eaten in Nigeria acceptable in 2020; Ogori et
nutrients and taste when grains are al., 2021)
partially substituted with fonio.
Miyan Touse When added to a traditional Nigerian (Helen &
or pumpkin soup, fonio increases nutrients. Ihekanu, 2009)
soup
Flour Fonio flours are gluten free and have (C. e. Chinma et
higher micro-nutrients than wheat al., 2015; Deriu
flours, and comparable functional et al., 2022; V.
properties with additions of 4% A. Jideani et al.,
carboxymethylcellulose (CMC) or 3% 2007;
xanthum gum. Compared to other Onyenweaku et
flours like maize, rice, sorghum, and al., 2021)
millet, fonio has higher water
absorption index, swelling power,
lower water solubility, and strong gel
properties.
19
Table 3.2 (cont’d)
Flour additive Fonio may also be added or combined
with other flours to increase nutrient
content of the flour. Many
combinations have been tried with
various levels of acceptability
(Supplemental Table__).
Malted Flour Malting grains increased the nutrients (J. A. Ayo et al.,
of the flour and functional properties 2019)
but had negative effects on taste and
other sensory properties according to
a panel.
Oil Fonio is low yielding for oil production (Ladan et al.,
but has high concentration of poly 2018)
unsaturated fatty acids and Vitamins A
and E.
Food Gums Fonio starch is suitable as a gum in (Arueya &
foods when the starch is modified by Oyewale, 2015;
succinylation. Olubusoye et al.,
2021)
Medicinal Fonio starch is suitable as an excipient (Akin-Ajani et al.,
tablets for medicinal tablets. Specific 2014; Emeje et
modifications such as acetylation, acid al., 2012; J. K.
hydrolysis, pregelantinized, or Muazu et al.,
carboxymethylation may improve 2009; Odeku,
functional properties. 2013; Omoteso
et al., 2019)
20
Table 3.2 (cont’d)
Animal Grain A 100% diet of fonio grains had a (Azi et al., 2021;
Feed positive or no effect on animal growth Fagbenro et al.,
and nutrient use compared to a maize 2000; Nwanna et
diet in fish, poultry, and rabbits. In egg al., 2003; Oke et
laying hens, 12.5% of the diet may be al., 2016; Sudik
replaced by fonio with no effect on egg et al., 2019; C.
production. Ukim et al.,
2012; C. I. Ukim
et al., 2013;
Wade & Audu,
2005)
Hay/Forage Known to feed cattle. Quality or yield is (Cruz et al.,
unknown. 2016)
Silage The fermentation process increases (Akinfemi, 2012)
crude protein and digestibility for
cattle.
Building OPC Fonio husk ash is an effective (E. E. Ndububa
Materials alternative Ordinary Portland Cement et al., 2016)
(OPC) in concrete.
Drywall Fonio husk combined with an adhesive (E. Ndububa et
liquid resin can be used as a particle al., 2015)
board, suitable for interior wall and
ceiling materials
Fuel Briquette Fonio husk may be turned into a (Bisu et al.,
briquette for use as fuel with good 2017)
density characteristics and ignition, but
poor burning rate.
Industry Enzymes for Fonio enzymes could be used for (Mulak &
fermentation brewing or ethanol production since Ogbonna, 2015;
the enzymatic activity was superior Nzelibe et al.,
compared to three other microbial 2000)
enzymes.
21
Table 3.2 (cont’d)
Silica Fonio husk ash was found to be an (Shamle et al.,
efficient alternative source of silica, 2014)
having higher yield and less mineral
contaminants compared to rice husk.
Biodegradable Fonio starch combined with glyceral (Alimi et al.,
packaging may be suitable for biodegradable 2021)
packaging materials.
Other Oxidized fonio starch may be useful as (Alimi &
a dispersant, emulsifying agent, Workneh, 2018;
adhesive, disintegrants, excipients, Isah et al., 2015)
flocculants, and other industrial uses.
Nutritional Quality
The micro and macro nutrient content of fonio has been well studied in the last 60 years
(Table 3.3) and is known to have high protein and micronutrient content (Table 3.4). Studies
testing fonio-based infant food in rats showed that fonio products had higher biological value
and protein efficiency ratios compared to maize products (Sudik et al., 2019). However other
studies found a positive or no effect of a fonio diet on growth and development in rats (Agbede
et al., 2019; Babarinde, Ebun, et al., 2020). A 10-20% fonio diet in rats also reduced blood
glucose levels (Osibemhe et al., 2020) potentially due to increasing insulin secretion (D. M.
Adams & Yakubu, 2020), low glycemic index (Robet et al., 2021), or alpha-amylase inhibitory
activity (Okonji et al., 2014). Nutritional studies in mice show benefits of a fonio diet and
consumers are known to eat fonio for its health benefits (Fogny-Fanou et al., 2010a; Kanupriya
& Dhiman, 2021), but there is little research determining the benefits of fonio on human health.
Fonio grains may be processed into a gluten free flour, and nutrients may be further
increased by creating flour mixtures or cooking method. Compared to common flours such as
wheat and oat, fonio flour had higher concentration of micro-nutrients (Onyenweaku et al.,
2021). However, like many other gluten free flours, fonio was not a source of folic acid,
representing a nutrient deficiency compared to wheat flours (Jamieson et al., 2020). Fonio flour
may be enhanced in protein and consumer acceptability by adding legume flour or other flours
such as sesame, plantain, or carrot (Supplemental Table 2). Malting fonio grains before grinding
for flour increased nutritional and functional properties, but had negative effects on taste and
22
sensory properties (J. A. Ayo et al., 2019). Extrusion cooking improved mineral availability,
protein dispersibility, and reduced antinutrients (Abiodun Victor et al., 2017; Anuonye et al.,
2007; Echendu et al., 2009; Ojokoh et al., 2015). However, cooking also has the potential to
decrease water soluble vitamin (Anuonye et al., 2007), phenolic, and antioxidant content (N’Dri
et al., 2013). At high cooking temperatures or long cooking times, the protein availability may
decrease as proteins are denatured (Chukwu, 2009). Processing and cooking methods are
known to affect fonio nutrient content, and more research is needed to understand optimal
methods to maintain bioavailability of nutrients.
While there are clear indications that fonio has potential as an alternative flour and has
nutritional properties that would be complementary to our diets, there are many production and
environmental factors that are not yet understood. For example, more research is needed on
genotype by environment interactions with the nutrient content of fonio grains. In a study
comparing micro and macro content across 104 samples and 3 environments in Mali, Barikmo
et al. (2004, 2007) found that micronutrient content (iron, niacin, thiamine) and macronutrient
content (protein, fiber, carbohydrates), was dependent on regions; suggesting that nutrient
quality is not consistent across regions. Several authors (Kawuyo et al., 2019; Kayodele et al.,
2019; T. K. Philip & Atiko, 2012) also found that moisture content of stored fonio grains before
dehulling effects physical properties like porosity, density, seed weight, sphericity, and specific
heat. It is not yet understood how moisture content may affect nutrient or antinutrient content of
fonio grains. Existing research on fonio has shown it has valuable nutritional properties, but
more research is needed to understand how those properties vary across varieties and
environments as well as the bioavailability of the nutrients.
Table 3.3: Count of studies analyzing fonio nutrients, sensory, and functional
properties for human consumption
Category Count
Major Nutrients: protein, carbohydrates, fat 18
Minor Nutrients: vitamins and minerals 11
Toxins or Anti-nutrients 6
Physical Characteristics:
grain size, 1000-grain weight, water absorption rate, 6
density, porosity, specific gravity
Fungi or Bacteria metabolite extractions 6
Flavonoids and Volatiles 7
23
Table 3.4: Nutritional properties of fonio grains
Details
Parameter Citation
(% per dry matter)
(Barikmo et al., 2004; V.
Clottey et al., 2006; De
Lumen et al., 1986; de
Protein Total 6-8%
Lumen et al., 1993;
Glew et al., 2013; Sadiq
et al., 2015)
Methionine 5-6% (De Lumen et al., 1986;
Lysine 2.6% de Lumen et al., 1993)
Fat 3-4% (Sadiq et al., 2015)
(Barikmo et al., 2004;
Carbohydrates 74-80%
Sadiq et al., 2015)
Fatty Acids 1.91% (Glew et al., 2013)
Ash 2.13-2.31 % (Sadiq et al., 2015)
Amylose 26.8% (Chukwu, 2009)
(Barikmo et al., 2004;
Fiber 0.80- 2.20%
Sadiq et al., 2015)
Copper 4.88 μg/g
Magnesium 1060 μg/g
Micronutrients Zinc 23 μg/g (Glew et al., 2013)
Calcium 172 μg/g
Sodium
0.02-0.03 μg/g
Potassium (Sadiq et al., 2015)
0.0054-0.00845 μg/g
Iron
0.00275-0.0011 μg/g
24
Table 3.4 (cont’d)
(Omaji et al., 2021;
Flavonoids 0.039- 0.05%
Sartelet et al., 1996)
Aldehydes, Heterocyclic,
Volatiles Identified: Alcohol, Ester, Furans, (Lasekan & Aina, 2003)
Ketone, Hydrocarbon
C-glycosyl (Boncompagni et al.,
Anti-Nutrients Low levels
flavones 2019)
Tannins Low levels
Ochratoxin A 0.00138-0.0239 μg/g (Makun et al., 2013)
Animal Uses
Fonio grains are economical and nutritious feed for rabbits, fish, and poultry. A 100%
diet of fonio grains had a positive or no effect on fish and poultry growth and nutrient use
compared to a maize diet (Fagbenro et al., 2000; Nwanna et al., 2003; Wade & Audu, 2005; C.
Ukim et al., 2012; C. I. Ukim et al., 2013, 2018). Female grower rabbits increased in growth
when 50% of their diet was fonio (Oke et al., 2016). In egg laying hens, 12.5% of the diet
replaced by fonio had no effect on egg production, and had a reduced cost in feed (Sudik et al.,
2019). For both fish and poultry studies, fonio grain was determined to be more economical than
maize due to increased profit indexes and less processing needed of fonio grains for feed
(Nwanna et al., 2003; C. I. Ukim et al., 2018, 2021). Ukim et al. (2021) reviewed the use of
fonio grain as feed for chickens and emphasized importance of fonio feed as normal feeds like
maize, sorghum, and millets are more expensive in northern Nigeria after the Covid-19
pandemic. However, it is unknown the economic impact on farmers when using fonio grains as
animal feed rather than for human consumption.
Fonio hay may also be used as feed for livestock as a straw, forage, or silage. It is
known that fonio is used to feed animals by grazing or cutting and storing (Cruz et al., 2016).
Akinfemi (2012) found that fonio straws averaged 6.28% crude protein and increased to 7.69%
crude protein when fermented for silage. In vitro digestibility also improved as fonio straw was
fermented for silage. Fonio may be suitable for silage and fonio straw may be improved through
this process. It is unknown how fonio forage compares to other common forages in nutrients
and in yield. It is also unknown if fonio is preferred by cattle. If grown as a forage crop, fonio
25
may be used in a variety of systems like a pasture or an annual forage mixture, but it is
unknown the performance of fonio in a mixture.
Material Uses
Fonio starch may be extracted and used in food and industrial applications
(Supplemental Table 3). Fonio starches at 15% concentration increased strength by 579%
compared to reference materials, making fonio starches applicable for making gluten free gel-
like foods (Deriu et al., 2022). Fonio starch was found to have similar angle of repose, cars
index, moisture conte, density, mean particle size to corn starch, and improved moisture
sorption and swelling power compared to corn starch (Musa et al., 2008). At concentration of
2% in weight, fonio starch compares to corn starch in flow rate, flow factor, tablet properties,
and may act as an alternative glidant or excipient for pharmaceutical use (J. Muazu et al.,
2010). These starches combined with glycerol and made into a film was found to have
applications for biodegradable packaging materials (Alimi et al., 2021). There are various uses
in industry, packaging, or food industry, but the applicability of these products is still not
understood. The cost of processing fonio starch has not been documented and it is unknown
how fonio starch products may compare economically to non-fonio products.
Typically, fonio’s husk is discarded as a waste product after the grain is cleaned, but
current research demonstrates other uses for the husk. When processed into ash, the husk is
an alternative raw source of silica used for a variety of application, having higher yield and less
mineral contaminants compared to rice husk (Shamle et al., 2014). The husk ash may also be
an effective alternative to Ordinary Portland Cement (OPC) in concrete (E. E. Ndububa et al.,
2016) or combined with an adhesive liquid resin and used as drywall (E. Ndububa et al., 2015).
The husk may also be turned into a briquette for use as fuel with good density characteristics
and ignition, but poor burning rate (Bisu et al., 2017). These uses to recycle a waste product
have not yet been economically analyzed for the financial impact to producers or the cost
compared to traditional materials.
3.5.3 Fonio Production
Overview
Fonio is grown across many different environments and cultures, but production remains
low with inconsistent levels of documentation. The distribution of studies in West Africa are
predominately in Nigeria and Mali (Figure 3.4). Most studies have come from Nigeria, totaling 16
studies which discuss production distribution, practices, farmer demographics, and cultural
26
perceptions of fonio. Many studies also come from Mali which detail the cultural perceptions of
fonio and compare production across environments. Although it is known that fonio is produced
in countries of Guinea, Burkina Faso, Ghana, and Ivory Coast, there are no documented
production practices or information on fonio farmers.
Although it is grown in many different agricultural systems, the production practices
remain relatively similar. As with many other crops in West Africa, fonio is produced primarily
without mechanization, requiring large amounts of labor to plant, weed, harvest, and process.
Labor is commonly split between men and women, with women completing the most labor-
intensive tasks. Fields are prepared by animal traction or hoe and then broadcast sown. Manual
weeding occurs 30-60 days after planting, primarily by women (Cruz et al., 2016; T. K. Philip &
Itodo, 2012). Inputs such as herbicides, pesticides, or fertilizers are rarely used in fonio
production. At harvest, men typically cut and stack sheaves to dry. Then it is threshed by
walking on the grains (women) or hitting the sheaves (men) to break off the grains. The grain is
gathered, sieved, and separated from the chaff by women. The process of dehulling the grain
and processing it for cooking is primarily done by women through iterations of pounding with
pestle and mortar and rinsing the grains (Cruz et al., 2016; Diop et al., 2018; T. K. Philip & Itodo,
2012).
As with many minor crops, women are heavily involved in the production and processing
of fonio. In Nigeria, 77% of fonio production is carried out by women (Istifanus & Agbo, 2016a).
In Senegal, fonio was ranked in importance by men who ranked millet, maize, and sorghum
above fonio and by women who ranked fonio higher than other cereals except for sorghum
(Diop et al., 2018). Processing of the fonio grain is also largely a task completed by women. In
Nigeria, 90% of processing labor is completed by women (T. K. Philip & Itodo, 2012) which is a
long and strenuous process. Cruz et al. (2016) reports five rounds of pounding and washing
totaling 43 minutes for the entire process and with an average of 32% yield loss from start to
finish, averaging 0.20 kg per hour per person. Of 16 studies on fonio production in Nigeria, only
two recognize gender as a factor in production. There is potential to expand gendered research
in fonio production and need for the inclusion of female voices in fonio research.
27
Figure 3.4: What is known about fonio production, practices, and farmers (left) and the
number of studies on these topics (right) throughout West Africa and the Dominican
Republic (Distribution of fonio production by Cruz et al., 2016 and FAO Stat)
Constraints to production and processing
Farmers have been surveyed in the countries of Nigeria, Senegal, Benin, Niger, and Mali
to determine major constraints to production. Across regions, the major constraint to growing
fonio is either low yields from small grain size or grain loss (Konate & Karambe, 2021; Popoola
et al., 2020) and/or high labor intensity of weeding, harvesting, and processing fonio (Diop et al.,
2018; Paraïso et al., 2016; T. K. Philip & Itodo, 2012; Sani et al., 2018) (Table 3.5). These
constraints contribute to young farmers and resource-scarce farmers being less likely to grow
fonio (Diop et al., 2018; Paraïso et al., 2016; T. K. Philip & Itodo, 2012; Sani et al., 2018). Diop
et al., (2018) also reports that fonio production is declining due to lack of seed but does not
explain the causes.
Various economic analyses provide more reasons for constraints on fonio production
which follow similar trends to other crop production in West Africa. These specific economic
analyses were completed in Nigeria. While findings may not be the same in other fonio
producing regions, they are likely indicative of general constraints in fonio production. Similar to
farmers’ listed constraints, the economic analyses determine that lack of mechanization is a
primary reason for inefficiency in fonio production (Kaka & Gindi, 2012). In Nigeria, in areas
28
where fonio is grown as a cash crop, production efficiency is around 80% (Duniya, 2013, 2015;
Duniya et al., 2015; Kaka & Gindi, 2012), and average profit index is 0.54 (Abdurrhman et al.,
2015). Inefficiencies are also attributed to issues that affect all farming practices in West Africa
such as lack of access to capital, credit, or improved seed varieties (Duniya, 2013; Duniya et al.,
2015; Kaka & Gindi, 2012). The cost of transactions such as marketing or the price of seed and
other inputs is another major constraint for Nigerian farmers (Duniya et al., 2015). There are
many opportunities to improve fonio production.
Table 3.5: Production constraints determined by farmers
Constraints were determined by multiple choice surveys or open-ended question interviews. An
asterisk indicates open ended questionnaires in data collection.
Country Constraints Citation
Harvest and weeding were the costliest and required the
most man-hours (T. K. Philip & Itodo,
Farmers most desired mechanization for dehulling and 2012)
harvest
Lack of improved seed, mechanization, and access to (Kaka & Gindi,
Nigeria credit to improve production 2012)
(Duniya, 2013,
High cost of labor for harvest and processing, high cost
2015; Duniya et al.,
of fertilizer and other inputs, inadequate capital
2015)*
Large amount of time and labor needed to weed and (Popoola et al.,
threshing grains, loss of grain during processing 2020)*
Decline in fonio production was attributed to lack of seed,
Senegal tedious processing, and disinterest of young farmers who (Diop et al., 2018)*
favor more productive crops or more profitable work
High labor requirement to produce fonio lead to less
young farmers or resource poor farmers not growing (Paraïso et al.,
Benin
fonio and devoting labor to more profitable income 2016)*
generation
High labor in post-harvest processing, lack of labor
Niger available since fonio harvest often coincides with major (Sani et al., 2018)*
grain crops
29
Table 3.5 (cont’d)
Low grain yields (<700 kg/ha) due to lack of improved (Konate & Karambe,
Mali
seed and optimal management 2021)*
Seed Systems
The seed system is an important aspect of production for sharing genetic materials and
dissemination of varieties. In Benin, the fonio seed system was recorded as being entirely
informal based on farmer exchange of seeds with each village, on average, maintaining one to
five varieties and individual households growing one to three varieties (Dansi, Adoukonou-
Sagbadja, & Vodouhe, 2010). Recording the seed system in more regions may identify
weaknesses or areas that could strengthen production. For example, in Mali, seed fairs and
community seed banks strengthened the seed system, increased production, and increased the
number of varieties grown (Sidibé et al., 2020). Seed systems will be increasingly important as
improved varieties are bred but may also benefit farmers now through access of varieties with
advantageous traits. The seed system is also crucial for conservation of genetic diversity.
Germination
Consistent germination is an important attribute of any crop and may be improved
through optimized biological and physical conditions. Hormone and photoperiod exposure were
evaluated for their effect on germination. Compared to controls with no added hormones,
germination rate increased with exposure to Thiourea (Idu et al., 2008) or presoaking seed in
0.2 ml Albit L-1 H2O (Anjorin & Salako, 2010). Germination decreased with exposure to 1-
naphthalene acetic acid (Idu et al., 2008). Additions of Albit+superhormai resulted in more
vigorous seedling growth, emergence, growth rate, and biomass compared to control with no
treatment (Anjorin & Salako, 2010). The effect of photoperiod on germination and seedling
development was tested with a single local variety from Nigeria. Germination rate was not
affected by photoperiod, but plant growth parameters (height, number of leaves, root length)
were optimal when germinated under a 12 hour photoperiod (Nyam, Angyu, et al., 2017). The
studied hormone or photoperiod treatment demonstrates that fonio germination may be
improved, but these studies must be expanded beyond a single variety. Other conditions for
germination may also provide valuable insights for planting or preparing fonio seeds. Moisture,
temperature, seed storage, and variety may play an important role in fonio germination.
30
Planting
Few studies have analyzed optimal planting methods. Traditionally, in most production
zones, fonio is planted at the onset of the rainy season (Adoukonou-Sagbadja et al., 2006; Cruz
et al., 2016). In southeast Senegal, an optimal planting time for early maturing varieties was
before July 15th which increased grain yield as much as 50-87% compared to later planting
dates (M. Gueye et al., 2015). More studies are needed to test optimal planting dates across
agroecosystems and varieties. Other factors which may affect planting time are variety,
agroecological zone, and photoperiod. While fonio is traditionally broadcast, drilling increased
plant growth parameters, straw yield, and grain yield (S. Dachi et al., 2017). Drill depth was not
reported but may be an important factor when planting because of small seed size. Drilling
would also require technology such as a tractor and a seed drill to practically implement.
Fertilizer
Fonio grain yields can range from as low as 210 kg/ha to as high as 1969 kg/ha
depending on production practices, varieties, and inputs. The first published research on
optimizing production for fonio was published in the 1980s on the use of fodder banks to
increase yields and was estimated that 40 kg/ha of nitrogen would be comparable to growing
fonio in fodder banks (Tarawali, 1988). Across three agroecological zones in Mali, an
observational study compared farmer practices of fertilizing and hand weeding, the highest
yields (1969 kg/ha) were observed with 100 kg/ha NPK (15-15-15) fertilizer and hand weeding
while the lowest yields were observed in zero input systems (Konate et al 2021). Four studies
support that increased fertilizer increases fonio grain and biomass yields (Table 3.6) with the
optimal range of nitrogen being between 15-30 kg/ha (Amekli, 2013; Bakare & Ochigbo, 2003;
S. Dachi et al., 2017; Gigou et al., 2009). Amekli (2013) demonstrated that for three fonio
landraces, the efficiency of grain production per unit of nitrogen increased and demonstrating
fonio has slight response to fertilization. These studies show that fonio yields benefit from low
level of fertilization, however, it is known that some fonio varieties are adapted to low fertility
soils and grown at the end of many rotations (Kanlindogbe et al., 2020). More research is
needed to understand the nutrient requirements of fonio and its niche in different soils across
environments and varieties.
31
Table 3.6: Summary and findings of fertilization studies.
Location Climate Soil Type Treatments Key Results Citation
Legon- Coastal Ferric 0, 30, 60 30 kg/ha NPK (Amekli,
Accra, Savanna Acrisol kgNPK/ha fertilizer increased 2013)
Ghana biomass and grain
yield 60% of three
landrace varieties
Birnin Sudan ferrugirious 0, 10, 20, 30 kg/ha N fertilizer (Bakare
Kebbi, Savanna tropical 30, 40, 50, increased tillers, &
Nigeria 60 kgN/ha plant height, and Ochigbo,
grain yields 2003)
Optimal rate of
return
Birnin Sudan ferrugirious 0, 30, 60 30 kg/ha N fertilizer (S. Dachi
Kebbi, Savanna tropical kgN/ha increased grain and et al.,
Nigeria biomass yield 2017)
Bareng & Tropical, Ferrasol 15 kgN/ha
Bordo, Savanna, Cambisol 0, 15, 30 increased grain and (Gigou et
Guinea Sahel Arenosol kgN/ha biomass yield al., 2009)
Cinzana, slightly to
Mali significantly across
locations and soils.
Rainfall was also a
limiting factor.
Drought Tolerance
Key physical and ecological characteristics of fonio demonstrate adaptations to drought
conditions. Documentation of fonio physical structure, and specifically epidermal structures have
been explored through microscopy (Irving & Jideani, 1997; Okanume et al., 2014), but these
structures were not related to drought tolerance until Dedeke (2021) found drought resistant
varieties with specialized cells to reduce transpiration and recover from oxidative stress.
Another drought resistance mechanism of fonio may be an association with microbes. Ndoye et
al. (2016) analyzed fonio’s response to arbuscular mycorrhizal fungi (AMF) and found fonio is
dependent on certain species of AMF and plant growth increases when associated with AMF.
32
These are the only known drought tolerant attributes and they have only been explored in a few
varieties. The metabolic, genetic, or environmental controls of these mechanisms are not well
understood. To apply these findings to breeding programs, the controls for these mechanisms
and the genetic markers need to be identified since not all varieties may exhibit drought
tolerance.
Pests and Pathogens
As with many crop species, weeds, pests, and pathogens can impact fonio growth and
post-harvest storage. In the field, yields may be decreased due to infection from striga (Terry,
1981) or other grass weeds (Umaru et al., 2010). These weeds are mainly controlled by hand
weeding during the first month of growth (Cruz et al., 2016). Stemborers were also found to be
the main insect pest of nine accessions in Nigeria along with several bird and rodent species
(Umaru et al., 2010). In Niger, grasshoppers reduce fonio yields and at times, have lead to
farmers abandoning fonio production (Sani et al., 2018). It is not recorded how insects, birds,
and rodent pests are controlled if at all. A leaf spot disease was identified to cause damage,
and can be controlled by a fungicide (Maji & Imolehim, 2013), or genetic resistance. Three
accessions (out of 23 studied) were identified to have resistance to leaf spot disease (Valerien
et al., 2020). After harvest, fonio grains may be susceptible to damage from T. confusum,
Aspergullus flavus, A. niger, and others with no suggested controls or prevention (M. T. Gueye
& Delobel, 1999; Umaru et al., 2010). Fonio is susceptible to many pathogens but has also
shown resistance and potential for use against other pathogens.
Fonio may serve a role in crop rotations to prevent or reduce the spread of disease to
other crops. For example, fonio reduced the presence of tobacco leaf curl virus when planted
right before tobacco (Deighton, 1940). A rotation of fonio grown in fields infected with M.
phaseolina also significantly decreased infection, benefiting cowpea yields compared to other
millet rotations (Ndiaye & Termorshuizen, 2008). The use of fonio in rotation for integrated pest
management have been tested, but more research is needed across environments, varieties,
and with other pathogens to determine the possibilities for fonio rotations.
Harvest
Harvesting fonio is a challenge due to shattering, lodging, and heterogeneous grain
maturation. Bakare (2005) demonstrated that optimal harvest time for a single fonio variety
grown in Badeggi, Nigeria was one week after physiological maturity and harvesting later
increases the potential of losing grain yield to shattering. Future studies may study optimal
33
harvest time based on variety and environmental conditions. Breeding efforts may also reduce
problems with grain shattering and lodging.
Processing
One of the major constraints to fonio production is the amount of labor to prepare the
grains for consumption, including dehulling and removing the bran. If this process were
mechanized, there may be significant benefit to fonio farmers, especially women. Different
researchers have evaluated modified sorghum (Marouze, 1994) and rice mills for use on fonio
(Marouzé et al., 2008). Other groups have developed dehullers (Idris et al., 2018; Tokan et al.,
2012) and continued to improve existing dehullers (Abdulrahman Abdulmalik et al., 2021; Ferre
et al., 2018; Kaankuka, 2015). Overall, the best recorded efficiency of a fonio dehuller is 69.9%
using a roller construction which increases total processing from 0.2 kg/hr by hand to 53kg/hr by
machine (Abdulrahman Abdulmalik et al., 2021). This mechanization will significantly decrease
the amount of time needed to process fonio. Dissemination of the technology will need careful
consideration to promote equitable access and benefit all fonio farmers.
Marketing
If the demand for fonio products increase, understanding the market will inform
production and breeding strategies. Selling strategies will benefit from documentation of
consumer preferences to direct the varieties grown and how the fonio is processed for market.
For example, in urban Mali, women will purchase higher quality fonio (based on color, milling,
and purity) at a price premium of 1-14% (Dury & Meuriot, 2010), meaning that farmers may be
able to get a higher price for certain attributes of fonio which are considered high quality.
Similarly, the greatest predictor of fonio consumption in urban Mali, was attitude toward fonio
(Fanou-Fogny et al., 2011) and two thirds of women surveyed had eaten fonio in the last month
and a majority (95%) of women believed it was a healthy food (Fogny-Fanou et al., 2010a).
Through surveys of producers and consumers, the perception of fonio has been recorded
through time and across regions (Table 3.7). Generally, it is known that fonio is eaten for health,
cultural, or food security reasons, but marketing strategies in specific areas would benefit from
greater documentation of consumer preferences and areas with the greatest demand for fonio
products.
34
Table 3.7: Cultural perception of fonio through time
Year Culture Perception Citation
Early Fouta-Djalon, (Cruz et al.,
Staple food
1900s Guinea 2016)
Dominican (Morales-payán
1940s Used as an aphrodisiac
Republic et al., 2002)
(Mazer & others,
1959 Mauritania Eaten when millet was scarce
1959)
Urban Mali, (Konkobo-
Appreciate and want to increase
2004 Guinea, Yameogo et al.,
consumption
Burkino Faso 2004)
2/3rds of women had eaten, and
Consumer (Fogny-Fanou et
2009 Urban Mali majority believed it’s good for
s and al., 2010a)
them
Producers
Sales increased during lean (Vall et al.,
2011 Rural Mali
periods 2011a)
(Kanupriya &
Dhiman, 2021;
2013 Senegal Used in traditional medicine
Koroch et al.,
2013)
Bauchi States, (Istifanus &
2016 Farmers hold “in high esteem”
Nigeria Agbo, 2016a)
Varied among ethnic groups and (Diop et al.,
2018 Senegal
genders 2018)
A secondary cereal that
1947 (Sudres, 1947)
contributes to soil degradation
Only a staple in "backward tribes
Research 1960 (Murdock, 1960)
of the northern Nigerian plateau"
ers
1977 A minor and lesser culture cereal (Harlan, 1977)
(T. Philip &
2006 A “lost crop” of Africa
Itodo, 2006)
An “economically important West (Mariac et al.,
2014
African crop species” 2014)
35
3.5.4 Fonio Breeding
Reproduction
Understanding the mode of reproduction is important for breeding and improvement of
crop species. Most Digitaria reproduce through apomixis, i.e. asexually without fertilization.
Similarly, fonio reproduces by apomixis with 1.7-2% outcrossing (Adoukonou-Sagbadja et al.,
2010; Barnaud et al., 2017). Because of the mode for reproduction, breeding by crossing parent
materials may be more challenging, but has not been attempted.
Genetics Tools
In the early 1900s, study of fonio genetics was limited to observational studies and
phenotypic data due to limited technology and general knowledge. As the field of genetics
expanded, biotechnology has expanded our understanding of fonio. Specific genetic tools and
their contribution to fonio research are outlined in table 3.8. A DNA extraction protocol and
Rapidly Detecting Genomic Polymorphisms (RAPD) analyses were optimized in 2005 to inform
future research (Kuta et al., 2005). SSR markers (Barnaud et al., 2012), ISSR markers
(Animasaun et al., 2017), and primers (Istifanus et al., 2018; Nyam, Kwon-Ndung, et al., 2017b;
Olodo et al., 2019) were identified as reference to the genome and potential site for marker
assisted breeding. Chloroplast genome sequences and reference transcriptomes have also
been identified for future use (Mariac et al., 2014; Sarah et al., 2017). The complete genome
was sequenced by Single Molecule Real Time Sequencing (SMRT)-cell technology in 2021
(Wang et al., 2021). Many of these tools have expanded our knowledge of fonio genetics and
origins but have not yet been applied in a breeding program.
Table 3.8: List of genetic tools and uses, review studies in bold.
Year Tool/Resource Major findings Citation
Rapidly detecting High genetic diversity of fonio
genomic with multiple domestication (Hitu et al.,
1997
polymorphisms (RAPD) and/or strong agroecological 1997)
analysis differentiation
36
Table 3.8 (cont’d)
DNA extraction
(Kuta et al.,
2005 protocol and RAPD Optimized for future use
2005)
analysis
(Adoukonou-
Flow cytometric Fonio genome size is stable Sagbadja,
2007
analysis across varieties Schubert, et
al., 2007)
Genetic distance- Genetic differentiation between (Adoukonou-
based clustering and fonio and D. iburu, three clusters Sagbadja,
2007
principal coordinate of fonio accessions related to Wagner, et al.,
analysis geographical origins 2007)
polymorphic simple
(Barnaud et
2012 sequence repeats Identified for future use
al., 2012)
(SSR) markers
Review of advances Specify genetic tools for better
(Barnaud et
2013 in plant genetic conservation of fonio genetic
al., 2013)
research resources
Protocol for complete
Barcoded libraries were (Mariac et al.,
2014 chloroplast genome
constructed for future use 2014)
sequencing in fonio
Summarized genetic controls
Review of genetic of shattering and hypothesize
(Patterson et
2016 mechanisms of shattering in fonio is controlled
al., 2016)
shattering by the homologous shattering
gene in rice, qSH1
37
Table 3.8 (cont’d)
Cluster analysis of ISSR markers
separated fonio and iburu into
two distinct clusters with an 0.71
Inter-Simple Sequence (Animasaun et
2017 similarity index, they hypothesize
Repeat (ISSR) markers al., 2017)
fonio and iburu have a common
ancestor but then speciated by
geographical isolation
Reference Reference database for future (Sarah et al.,
2017
transcriptomes use 2017)
Identified distinct genetic (Nyam, Kwon-
2017 differences between fonio and Ndung, et al.,
Microsatellite primer
two related species 2017b)
combinations designed
Three primer pairs successfully
and tested (Istifanus et al.,
2018 separated four fonio types based
2018)
on grain color
126 new primer pairs
Confirmed fonio is closer related (Olodo et al.,
2019 identified for SSR
to D. longiflora than D. iburu 2019)
markers
Fonio diversity is shaped by
Chromosome-scale
climate, geography, and
reference assembly
ethnolinguistics (Abrouk et al.,
2020 and re-sequencing of
Genes associated with shattering 2020)
183 accession of
and seed size show signs of
Digitaria
domestication
Fonio genome Tetraploid genome with
sequence with long- homoeologous duplications
(Wang et al.,
2021 read Single Molecule Diversity in Mali was distributed
2021)
Real Time Sequencing north to south along
(SMRT)-cell technology agroecosystems
38
Genetic Diversity
Fonio has high genetic diversity primarily because of geographic differentiation. In a
large study across five countries (Benin, Burkino Faso, Guinea, Mali, and Togo) and 122
varieties, three genetic groups were differentiated by geography and country of origin
(Adoukonou-Sagbadja, Wagner, et al., 2007). A chromosome reference assembly created with
183 cultivated and wild varieties of Digitaria found that fonio diversity has been shaped by
climate and geography (Abrouk et al., 2020). Sequencing of the fonio genome and further
genetic analysis of varieties from Mali, also suggest genetic differentiation by geography,
especially between the Sahel and grasslands of Mali (Wang et al., 2021). Fonio varietal
genetics are diverse, but also stable in genome size (Adoukonou-Sagbadja, Schubert, et al.,
2007). This diversity is well documented, and efforts have been made to conserve this essential
resource by the assemblage of diverse genetic materials. Collections have occurred since the
1970s and have coincided with documentation of variety attributes and farmer preferences
(Table 3.9).
Table 3.9: Record of variety collections including the country, year, and data
collected
# of
Country Year Accessions Other Data Collected Citation
Collected
(Bakhareva & Mukha,
Nigeria 1971 Yield
1971)
(Kwon-Ndung et al.,
Nigeria 1995 138 Farmer demographics
1998)
Farmer classification and
selection parameters:
(V. A. Clottey et al.,
Ghana 2000 11 maturity, ease of
2006)
processing, storage
qualities
(Adoukonou-Sagbadja,
Togo 2001 95 Village ethnic group
2004)
Origin, cooking qualities, (Adoukonou-Sagbadja
Togo 2008 42
growth cycle, color, size et al., 2008)
39
Table 3.9 (cont’d)
Multiple 2013 20 Sensory characteristics (Fliedel et al., 2013)
Variety names, uses,
Benin 2014 35 (Ballogou et al., 2014)
agronomic characteristics
Maturity, yield, plant
Benin 2016 20 (Sekloka et al., 2016)
architecture, plant height
Major use, qualities of
(Istifanus & Agbo,
Nigeria 2016 14 farmers, farmer opinion of
2016b)
fonio
Plant characteristics, 1000 (Nyam, Kwon-Ndung,
Nigeria 2017 30
seed weight et al., 2017a)
Root architecture and (Ogunkanmi et al.,
Multiple 2018 75
drought tolerance 2018)
Key Traits
These varietal collections also document key traits, allowing potential breeding materials
to be identified. The greatest agronomic differences between varieties were grain yield, time to
maturity, plant height, and panicle or leaf characteristics (S. N. Dachi & Gana, 2021; Nyam,
Angyu, et al., 2017; S. I. Saidou et al., 2014; Sekloka et al., 2016). Grain yield was found to be
correlated positively to number of spikelets (A. Aliero & Morakinyo, 2002), plant height (Nyam et
al., 2021), and time to maturity (Sekloka et al., 2016). Days to flowering was correlated to
photoperiod and variety (A. A. Aliero & Morakinyo, 2005). Drought tolerance was also found to
correlate with root architecture, specifically length of primary and lateral roots (Ogunkanmi et al.,
2018). The next steps for using these traits for varietal improvement will be to identify genetic
markers and controls for these traits.
Heritability
Very few studies have studied the heritability of agronomic fonio traits. Nyam et al.
(2021) found that across 10 varieties, number of spikelets per panicle, number of tillers per
plant, and 1000 seed weight had high heritability and may be used for selection to increase
grain yield. The study also found that across all traits, phenotypic variance was greater than
genotypic variance, indicating a strong environmental influence on variety performance (Nyam
et al., 2021). In Niger, a study that evaluated 67 accessions of fonio across two environments
40
found that differences in varietal growth parameters were greatly influenced by environment and
genetic, environment interactions (S. Saidou, 2017). They suggest that in future breeding
efforts, fonio must be tested across multiple environments.
Mutagenesis
Because of the challenge of crossing parent materials, mutagenesis may be a key
method to create more genetic diversity in a breeding program. Six studies have identified
methods for creating fonio mutants with improved traits either chemically, biologically, or
physically (Table 3.10).
Table 3.10: List of mutagenesis studies and their major findings, reviews in bold
Tool for
Major findings Citation
mutagenesis
Mutagenesis is useful for creating
genetic diversity, polygenic (Sarker et al.,
Review
variation, and inducing male 1993)
sterility
4 hr of exposure to 0.1M Nitrous acid (Animasaun,
Nitrous acid resulted in improved growth and yield Mustapha, et al.,
Chemical of mutants 2014)
Produced taller, more leafy mutants (Nura et al.,
Colchicine
with more tillers and grains per spike 2017)
Agrobacterium- Genes were successfully transferred
(Ntui et al.,
Biological mediated genetic from the agrobacterium to two
2017)
transformation varieties of fonio
80Gy improved germination, plant
(Animasaun,
Gamma growth parameters (height, tillers,
Morakinyo, et
irradiation leaves), early maturity, and 100 grain
al., 2014)
weight
Physical Gamma 100Gy induced the most (Nura et al.,
irradiation advantageous agronomic traits 2021)
60 min of exposure improved
Alpha-spin (Goler & Kwon-
performance and three varieties were
nanoparticles Ndung, 2020)
identified as the best performing
41
Farmer Selection Criteria
Improvement of fonio varieties is needed especially related to agronomic characteristics.
However, it is important to note that West African farmers have been selecting for specific traits
for thousands of years and may provide insights to prioritize breeding objectives. Based on
farmer surveys, farmers generally select based on rate of maturation, agronomic characteristics,
and grain characteristics such as ease of processing, storing, cooking, color, or size (Table
3.11). The specific trait farmers select for like early maturation vs. late maturation depends on
the region and the farmers’ intended use for the crop. For example, in Benin, earlier maturing
varieties were preferred since the crop could fill a lean period before the main harvest (Dansi,
Adoukonou-Sagbadja, & Vodouhè, 2010), but in areas where fonio is produced as a cash crop,
longer maturing varieties with higher grain yields may be preferred. These preferences are also
gendered with women preferring varieties with an easily removed hull, good taste, or other
cooking qualities. Men preferred higher yielding fonio varieties with greater drought tolerance
and less lodging for ease of harvest. To ensure acceptability and adoption of improved varieties,
farmers may aid in the selection process. This is also an opportunity to support women by
inclusion of their preferences.
Table 3.11: Farmer* criteria for varietal selection
Country Criteria for Selection or Preference Citation
Early maturation, ease of processing (softer (V. A. Clottey et al.,
Ghana
hull), and storage qualities (unknown specifics) 2006)
Early maturation, cooking qualities, and grain (Adoukonou-Sagbadja
Togo
lighter in color with larger in size et al., 2008)
Early maturation, cooking qualities, ease of
(Dansi, Adoukonou-
processing (softer hull), productivity, ease of
Benin Sagbadja, & Vodouhè,
harvest with decreased lodging, larger grain
2010)
size, storage, and drought tolerance**
cooking qualities, ease of processing, and
Benin (Ballogou et al., 2014)
agronomic traits
*gender not specified
** indicated different preferences by gender
42
3.6 Discussion
As summarized above, there are many gaps in our knowledge of fonio across disciplines
which may be analyzed through an agroecological framework and applied within fonio’s current
system as a food security crop or in new systems. Specifically, within the agroecological
framework, there are key topics that can be addressed in future fonio research. Diversity may
be addressed both in the functional diversity of agroecosystems and the role fonio may play in
increasing functional diversity and in understanding the diverse cultural value of fonio.
Efficiencies may also be improved in fonio production through breeding and mechanization and
through recycling of waste products such as the husk and the stover. Across many disciplines,
the recognition of human and social values will be essential to inform breeding objectives and
production improvements not only across diverse cultures but also across genders and
socioeconomic groups. Future research and development of new markets will rely on
governance mechanisms which are “transparent, accountable, and inclusive”, summarized as
responsible governance (FAO, 2018).
Major Gaps in Current Production Systems
Fonio is produced across West Africa, but more documentation is needed to understand
the diversity of management strategies, varieties used, and cultural value of fonio. While these
aspects have been recorded in a few locations (Table 3.7), our overall understanding of fonio
may benefit from greater documentation. This connects back to the agroecological element of
understanding the culture and food traditions of fonio, helping to honor the stewards and protect
the cultural value of fonio for future generations. Similarly, there are recordings of varieties used
and specific attributes that have been made in a few regions (Table 3.9), while others may still
be undocumented. The current systems may benefit from wide exchange of varieties which may
have advantageous traits like drought tolerance and yield. Strengthening the informal seed
system for other minor crops has been shown to be effective for conservation of biodiversity and
increasing food security (Gill et al., 2013). Likewise, the trade of fonio varieties with
advantageous traits may enhance the ability for fonio as a food security crop and increase
diversity and resiliency of these cropping systems.
Breeding is another major development that may improve food security, but better
defined breeding objectives are needed with farmer preferences included. We hypothesized that
breeding programs would improve production and found tools that have been developed for
breeding fonio. The gap in this literature is the application and result of these tools through
breeding programs. Initial breeding objectives should focus on improved agronomic aspects to
43
increase production and decrease labor such as reduction of shattering and increased grain
size. Greater documentation of consumer grain preferences and the intended culinary use
(porridge, flour, or brewing) will also inform breeding objectives. Interdisciplinary teams that
include breeders, agronomists and gender specialists need to document knowledge on traits for
specific end-uses and environments, taking into consideration selecting for specific traits such
as reduced seed shattering or larger grain may impact a desirable end-use trait (Diallo et al.,
2018; Weltzien et al., 2019). This is an opportunity to create knowledge with fonio farmers that
could serve to conserve traditional knowledge and genetic resources.
Another major area of improvement needed for food security is mechanization of grain
processing which could increase capacity and reduce labor requirements. Women are heavily
involved in fonio production, especially processing the grains for cooking, thus mechanization of
processing has the potential to empower or disempower women. Mechanized tasks often
become male-dominant even when the task was originally done by women (Fischer et al.,
2018). Women’s labor burden may be reduced by men taking over mechanized processing of
grains, thus allowing for more choice in how to spend time. Increased grain processing may
allow farmers to sell fonio instead of saving for home consumption. This could impact food and
nutritional security since fonio is often used during lean periods and beneficial for dietary
diversity (Conti et al., 2019). Increased income from fonio sales may or may not benefit women
depending on the use of the income and the extent to which they have decision-making power
in the use of income (Ganle et al., 2015; Mudege et al., 2015). Women and other marginalized
farmers may also be further disadvantaged with a potential loss of a valuable food source if they
are unable to access the mechanized technology (Eidt et al., 2020). While mechanization would
substantially increase production potential, it may also become a disadvantage for traditional
stewards of the crop and reduce women’s control over a household food staple and nutritional
security. When implementing these technologies, it will be essential to establish community
groups that agree on fair and equitable use with mechanisms for resolving challenges (Fischer
et al., 2018), contributing specifically to the agroecological element of responsible governance.
Major Gaps in Knowledge of Fonio in Novel Systems
When expanding the use of fonio in new cropping systems, a major gap in our
knowledge is the effect of variety and environment on specific traits. Much of the research
exploring fonio management has only been done on a single variety and environment. Thus, it is
unknown if resilient traits are only present in certain varieties or environments which will become
44
even more important if fonio production expands beyond the West African context. There is
already evidence that nutrient content is dependent on variety or environment (Barikmo et al.
2004, 2007), but the effects have not been studied enough to understand the relationship
between nutrition and growth environment. Further research is needed on genotype by
environment (GxE) interactions and the effect on traits, yield, and nutritional quality. Globally
testing varieties of quinoa helped the expansion and continued research, as its potential uses
and benefits were understood (Bazile, Jacobsen, et al., 2016). Likewise, global variety testing
may help expand fonio production and contribute to the resiliency of cropping systems globally.
Research is needed to identify how fonio may benefit global diversity and resilience. The
greatest benefits from increased crop diversity often are from functional diversity, where
different crop species use different niches and available resources are more fully utilized (Lin,
2011; Song et al., 2014). Studies have shown that increasing plant biodiversity with functional
diverse groups such as forbs, legumes, and grasses help build soil fertility compared to
monocultures (Furey & Tilman, 2021). However, it is not understood the specific niches that
fonio uses in the soil or how it impacts microbial diversity. Likewise, it is not known how fonio
may react to interspecific competition which would be important for utilizing fonio in a mixture.
Fonio is also known to have reliance on microbes which aid in drought and low nutrient
conditions (Ndoye et al. 2016), but it is not understood how these interactions may change as
fonio is grown on different continents with potentially new microbes. To fully utilize fonio globally
and improve diversity and resilience, it first must be understood how growth and adaptations
translate to new cropping systems.
Research in fonio breeding and mechanization may also improve fonio production in
new locations and management systems. Breeding objectives would be dependent on the
intended end use of the crop as a grain, forage, or other use. Major breeding objectives for
growing fonio in mechanized systems should focus on crop properties that allow mechanization
such as upright growth habit, non-shattering, and higher yielding. For global production,
mechanization is essential to increase production and reduce labor requirements but will require
breeding programs and optimized management strategies according to the end use.
Increased global demand for fonio products may benefit or harm fonio farmers and
requires responsible governance to ensure equity. Like the boom and bust trends observed in
tef and quinoa (Andreotti et al., 2022), the initial increase in demand for fonio will increase
production and income-generation for farmers. Research will likely improve production and
potentially increase production outside of West Africa but may not benefit fonio producers in the
long term. If industrialized countries or private companies take over production, then the price of
45
fonio in West Africa may rapidly decrease and make imported fonio products cheaper than
locally grown (Andreotti et al., 2022). The original growers and stewards of fonio may then be
forced to buy fonio and no longer profit from fonio sales. In Ethiopia, tef exports were banned in
an effort to maintain national food security and secure their genetic resources, but this was
ineffective to prevent the boom and bust cycle (Crymes, 2015). There was greater success in
quinoa protection in Andean countries with the creation of international research groups,
farmers groups, NGOs, and the increased production of highly valued varieties that helped to
maintain benefits to local producers. However, the results have varied and still result in
disempowerment of some farmers who grew quinoa traditionally (Drobnack & Isaacs, in review).
Increased non-food fonio uses may further impact food security and traditional fonio
uses. For example, selling crops as biofuels may improve financial security of resource-poor
farmers and thus decrease vulnerability to climate and price shocks (Ewing & Msangi, 2009).
There are still tradeoffs to growing crops for food versus nonfood uses. Many smallholder
farmers grow crops for home consumption. Increased purchasing power may not always
increase food security if the additional income is not used to purchase food. Additionally, if fonio
is grown for non-food uses, then fonio processed as grain may become even more expensive,
leading to less consumption of fonio and loss of cultural value (Ewing & Msangi, 2009).
Therefore, great care must be taken in creation of fonio markets to support the cultural value
and stewardship of fonio producers.
3.7 Conclusions
The scoping review sought to summarize fonio research across disciplines with an
emphasis on production and the implications of research using an agroecology lens. Research
on fonio has steadily increased in the last 50 years with a great emphasis on nutrients and
culinary uses. Many novel uses have been identified, and to support the increased use of fonio
as a food security crop or as a novel crop globally, more research is needed. Although there are
many knowledge gaps, we focused on fonio production which may benefit from greater
understanding of management strategies particularly across varieties and environments.
Mechanization of grain processing may also increase capacity for fonio production and reduce
labor requirements for women. Breeding programs would also greatly benefit fonio production
by decreasing yield loss from shattering and lodging. Each of these may benefit from
partnership with farmers who would provide valuable knowledge. As production and research
46
increase, care must be given to ensure women are empowered, and producers are protected
from the “boom and bust” cycle of minor crop development.
3.8 Supplemental Materials
Supplemental Table 1: Descriptive summaries documenting information on fonio
Year Title Scope of Publication Citation
1955 The minor Summary of descriptive information given (Porters,
cereals of the on D. exilis including: history, synonyms, 1955)
genus Digitaria in botanical description, germination, seedling
Africa and development, vernacular names, crop areas,
Europe varieties, cultural methods, harvest, yield
and uses. Information is also given on D.
sanguinalis and D. iburua.
1960 Staple Summary of major crops throughout Africa. (Murdock,
subsistence Includes the distribution of fonio in West 1960)
crops of Africa Africa and indicates it is likely one of the
oldest cultivated grains and is often replaced
by sorghum and millet. Denigrates fonio as
eaten only by uncivilized tribes in Nigeria
1974 West African Summary of many crops from Africa. (Lewicki,
Food in the Records three kinds of fonio grown in West 1974)
Middle Ages Africa: animal fonio or guinea millet
(Brachiaria deflexa), true fonio (Digitaria
exilis), and black fonio (Digitaria iburu) with
the distribution of each
1977 The origins of Summary and history of cereals both major (Harlan,
cereal agriculture and minor cereals with fonio briefly 1977)
in the Old World described.
1996 Lost crops of Summary of crop production areas and (Council &
Africa: Volume 1: environmental requirements, uses, others, 1996)
Grains nutritional value, agronomy, harvesting and
handling, limitations, and prospects with
fonio included in one chapter
47
Table 1 (cont’d)
2002 Digitaria exilis as Summary of fonio, called Funde, in the (Morales-
a Crop in the Dominican Republic. Brought in the early payán et al.,
Dominican 1500s, fonio survived mostly as a weed until 2002)
Republic cultivation began in the 1940s as a grain and
as a forage.
2006 Acha Summary of Digitaria exilis and Digitaria (T. Philip &
(Digitaria spp.) a iburua nomenclature, history, and Itodo, 2006)
"rediscovered" importance.
indigenous crop
of West Africa.
2012 Digitaria exilis Summary of potential of fonio diets, (A. I. O.
(acha/fonio), wellness, economic status improvement, and Jideani,
Digitaria iburua role in food as a traditional food. Call for 2012)
(iburu/fonio) and greater collaboration and development to
Eluesine increase the use and benefits from these
coracana crops.
(tamba/finger
millet) Non-
conventional
cereal grains with
potentials
2015 Teff & Fonio – Summary of two African crops- their (Small, 2015)
Africa’s cultivation and uses.
sustainable
cereals
2016 Drought Summary of drought tolerance and (Tadele,
Adaptation in strategies in several millets, including fonio. 2016)
Millets
48
Table 1 (cont’d)
2016 Vernacular The paper uses the patterns discerned in (Blench,
names for African vernacular names to explore their history. 2016)
millets and other
minor cereals
and their
significance for
agricultural
history
2016 Fonio, an African Summarizes the practices, yields, and (Cruz et al.,
cereal culture associated with fonio cultivation. 2016)
Records developments in mechanization.
Supplemental Table 2: Flour blends tested for acceptability and/or benefits
Most Acceptable
Flour Components Benefits Citation
Composition
Flavor, consumer
acceptability (McWatters et al.,
Fonio, Wheat 50:50
Comparable to 100% 2003)
wheat bread
Highest baking
potential, dough (Drábková et al.,
5:95
machinability, 2017)
nutritional benefits
Nutrients, sensory,
Fonio, Wheat, (Orisa & Udofia,
23:50:25:2 consumer
Cowpea, Moringa 2019)
acceptability
Fonio, Wheat, Protein, fiber,
70:15:15 (Nanyen, 2016)
Mungbean minerals, fat, vitamins
49
Table 2 (cont’d)
Consumer
Fonio, Wheat, Kidney 00:75:25 or (Inyang et al.,
acceptability, fiber,
bean 25:50:25 2018)
nutritional value
Fonio, Wheat, Sensory, consumer (Adeyanju et al.,
50:20:30
Pigeonpea acceptability 2018)
Fonio, Wheat, Pasting properties, (Olagunju et al.,
Pigeonpea protein 2020)
Consumer
Fonio, Wheat, (Emelike et al.,
40:30:30 acceptability, protein,
Cashew kernel 2020)
vitamins, minerals
Fonio and Bambara
Consumer (C. E. Chinma et
nut sourdough, 10:90
acceptability al., 2016)
Wheat
Fonio, Wheat, Protein, fiber, ash, (Olagunju et al.,
10:70:20
Amaranth lower glycemic index 2021)
Complementary (Olapade & Aworj,
Fonio, Cowpea
proteins 2012)
Consumer preferred,
Fonio, Cowpea, decrease wheat (Inyang &
56:20:24
Banana importation, increased Nwabueze, 2020)
nutritional composition
Comparable to 100% (V. A. Jideani et
Fonio, CMC 96:4
wheat bread al., 2007)
Fonio, Potato starch, Protein, nutrients, (V. Jideani et al.,
73:20:2:5
Yeast, Soybean sensory 2008)
Fonio, malted Gluten and sugar free,
Bambara nut, Date 60:10:30 consumer (Agu et al., 2020)
palm acceptability
Fonio, Bambara nut,
80:10:10 Protein, fiber, nutrients (Agu et al., 2014)
Plantain
50
Table 2 (cont’d)
Fonio, Bambara nut, Protein, b-carotene,
(Okafor, 2016)
Carrot minerals
Nutrients, functional
Fonio, Soybean, 70:15:10:5 or (Ukeyima et al.,
properties, sensory
Carrot, Milk flavor 60:25:10:5 2019)
characteristics
Gluten free, consumer
acceptability,
Fonio, Bambara nut, (C. e. Chinma et
75:25:3 comparable to wheat
Xanthum Gum al., 2015)
bread in storage and
flavor traits
Fonio, Soybean, Chemical and
70:20:10 (Ikujenlola, 2008)
Sesame functional properties
Protein, fat, energy,
Fonio, Soybean, (Combo et al.,
but low in
Mango 2020)
micronutrients
Fiber, minerals, fats,
Fonio, Soybean husk 97.5 : 2.5 proteins, consumer (Adekunle, 2018)
acceptability, texture
(Olagunju et al.,
Fonio, Pigeon Pea 70:30 Protein, antinutrients
2018)
Protein, ash, amino (Babarinde,
Fonio, Pigeon Pea acids, vitamins, Adeyanju, et al.,
sensory analysis 2020)
Protein, fiber, fats,
Fonio, Moringa 88:12 (Raji et al., 2018)
vitamins, minerals
Phytochemicals,
sensory properties, (J. Ayo, Ayo, et al.,
Fonio, Orange Peel 95:5
consumer 2018)
acceptability
Fiber, fat, crispiness, (Adekunle et al.,
Fonio, Tigernut
texture 2018)
51
Table 2 (cont’d)
Fonio, Tigernut, Protein, amino acids,
50:45:5 (Adesida, 2020)
Sorrel Seed fatty acids, minerals
Sensory, consumer (J. Ayo, Ojo, et al.,
Fonio, Mushroom 95:5
acceptability 2018)
Protein, ash, fat, fiber,
(J. A. Ayo et al.,
Fonio, guava 95:5 consumer
2020)
acceptability
Fonio, Sandpaper Better hypertensive-
96:4 (Oboh et al., 2021)
leaf diabetic properties
Protein, Fiber,
Fonio, Mango kernel, (Olorunfemi et al.,
Flavonoid, phenol,
Soy Cake 2022)
less antinutritients
Comparable to 100% (J. Ayo et al.,
Fonio, Banana Peel 75:25
wheat bread 2022)
Minerals, antinutrients,
(Obizoba &
Fonio, Baobab 70:30 protein
Anyika, 1994)
CMC= carboxymethylcellulose
Supplemental Table 3: List of starch modifications
Modification Properties Potential uses Citation
Reduced physiochemical
properties:
Foaming, amylose content, (Emeje et al.,
Acetylation
true density, bulk density, 2012)
water absorption capacity,
ash content, and pH
Improved physiochemical Compressible (Akin-Ajani et al.,
Acid hydrolysis
and compaction properties excipients 2014)
52
Table 3 (cont’d)
Functional and
Gums in food (Arueya &
Succinylation morphological properties
application Oyewale, 2015)
identified
(Olubusoye et al.,
2021)
Dispersants,
emulsifying
agent, surface
sizing,
Oxidization Improved stability (Isah et al., 2015)
adhesive,
disintegrants,
excipients, and
flocculants
Improved disintegrant
properties Paracetamol (Akin-Ajani et al.,
Acid
Comparable to corn starch tablet material 2014)
disintegrants
Improved functional (Alimi & Workneh,
Citric acid Industrial uses
properties 2018)
Carboxymethylation Improved drug release Pharmaceutical (Omoteso et al.,
Pregelatinised properties uses 2019)
Low onset of plastic
deformation, increased
crushing strength, Paracetamol (J. K. Muazu et al.,
Pregelatinised
disintegration, binder tablet material 2009)
concentration, and
dissolution time
53
4. GROWING FONIO IN MICHIGAN: THE EFFECT OF SEEDING AND FERTILIZER
RATES ON FONIO GRAIN YIELD, BIOMASS PRODUCTION, AND FORAGE
QUALITY
4.1 Abstract
Crop diversity is essential in supporting many ecosystems’ services including the resilience of
agroecosystems. As climate change increases stressors, resiliency is essential for food and
nutritional security. Greater utilization of minor crops can benefit U.S. cropping systems by
building resiliency and filling specific niches. Fonio is a small grain traditionally grown in West
Africa with great potential as a climate smart crop. Lack of global recognition has limited crop
improvements and optimization of production practices. To test fonio in a new environment and
explore its uses, fonio was grown in Western Michigan in 2021 and 2022. Two trials were
conducted to test seeding rates and fertilizer rates. Total above ground biomass, grain yield,
growth parameters, and forage potential were measured. The results highlight several
challenges associated with fonio production including lack of flowering, weed management, and
grain processing. Lower seeding rates and low fertilization increased biomass yields but had
little to no effect on grain yields. Relative forage quality was high (130-150) when cut at booting,
but yields were low (2249-5585 kg/ha) in comparison to other forages. Fonio may be cut several
times for forage, but quality is reduced with subsequent cuttings. Research is needed to
understand how variety and environments affect fertilizer response and forage quality, and to
continue evaluating fonio within U.S. cropping systems.
4.2 Introduction
Diversity in agricultural systems is essential for supporting ecosystem services.
Increasing functional diversity is known to improve long term community productivity and
therefore soil health such as increased nitrogen, potassium, calcium, magnesium, and carbon
(Dybzinski et al., 2008; Furey & Tilman, 2021). Addition of a single crop in rotation can increase
soil carbon, nitrogen pools, and soil microbial biomass (McDaniel et al., 2014). Proper
management of diversity can also prevent pest and disease outbreaks, reducing pesticide
application. For example, an infestation of fall army worm was reduced 70-96% by increasing
the number cover crops planted in a mixture (Meagher et al., 2022). On a landscape scale,
diversity increases the density and species richness of pollinating insects potentially increasing
yield over time (Aguilera et al., 2020; Raderschall et al., 2021). In integrated crop-livestock
systems, greater forage diversity can improve yields and economic return (Sanderson et al.,
2013) while providing a more diverse diet for livestock.
Crop diversity has additional benefits for resilience and stability as stress on agricultural
systems increase due to climate change. Climate change is impacting temperature and
precipitation throughout the U.S. by making weather events more extreme like long droughts,
more intense storms, and hotter summer temperatures. As climatic variability increases,
diversity in species or variety may improve yield stability across environments (Bybee-Finley &
54
Ryan, 2018; Reiss & Drinkwater, 2018). Increasing the number of crops grown regionally or on
a farm scale will decrease the likelihood of complete crop loss through the insurance hypothesis
(Lin, 2011). Resilience of agroecosystems to maintain functionality during or after a disturbance
increases with improved ecosystem function. Functional diversity improves ecosystem function
by increasing soil fertility and water conservation (Altieri, 2004). Climatic stressors will also likely
increase biotic stress from pests and pathogens, increasing the importance of pest
management and the use of crop diversity to prevent outbreaks.
Minor crops offer an opportunity to increase biodiversity and fill niches which may not yet
be filled by major crops. Many of these minor crops are under-researched and are used in the
U.S. as cover crops, niche grains, or forage crops. Crabgrass (Digitaria sanguilis) is often
viewed as a weed in the United States but is also a drought tolerant forage crop that can
increase the water use efficiency of U.S. pastures (Gelley et al., 2020). Cowpea and sun hemp
used as cover crops in southern Texas and Florida may reduce the pressure of fall army worms
(Meagher et al., 2022). Tef, a small grain from Ethiopia, is grown in the U.S. as a grain for
human consumption or as a high quality forage crop. As a forage, tef is grown during the hottest
and driest summer months with high quality suitable for horse feed (Fitting Teff into the Horse
Diet, n.d.). As a grain, tef can be processed into a gluten free flour which will aid those with
celiac’s disease or gluten intolerance (Teff Is a Healthy Wheat Alternative, n.d.). Fonio is
another small grain from West Africa that is drought tolerant and may benefit the resilience of
U.S. agroecosystems.
In West Africa, farmers and crop stewards have been cultivating a small grain, fonio
(Digitaria spp.), for thousands of years. This tiny grain is a warm season grass of high value to
farmers in the region as a nutritious crop for low fertility sites with drought tolerant properties
((Fogny-Fanou et al., 2010a; Vall et al., 2011b). Often, it is produced at the end of many crop
rotations without external inputs (Kanlindogbe et al., 2020). In U.S. systems, fonio would likely
need very few external inputs and may be grown during drought or as a second crop. Fonio is
also known to have a negative effect on the lifecycle of certain infectious fungus species
(Kanlindogbe et al., 2020; Ndiaye & Termorshuizen, 2008), making it a possible cover or
rotation crop to prevent pest outbreaks. With its preserved genetic diversity, there may be other
unidentified traits. Just in Togo, 42 different varieties of fonio were characterized with a variety
of agronomic attributes (Adoukonou-Sagbadja et al., 2008).
In U.S. systems, fonio may also be used as a niche grain for brewing or eating. It is
growing in popularity in restaurants in the United States and there are several fonio brands
currently being imported to the U.S. A Brooklyn brewery is currently selling beer made from
55
fonio grains (Swierk, 2022) and a chef in New York is developing recipes for Western
consumers (Hunt, 2020). As with quinoa and tef, fonio may find a place is U.S. food systems as
a healthy alternative to major grains.
Fonio may have an economic advantage as a dual purpose crop. It is known that fonio
hay is used to feed animals by grazing or cutting and storing (Cruz et al., 2016). Akinfemi (2012)
found that fonio straws averaged 6.28% crude protein and increased to 7.69% crude protein
when fermented for silage. In vitro digestibility also improved as fonio straw was fermented for
silage. However, it is unknown how fonio forage, straw, and silage compares in nutrients or in
yield to other common forages.
The lack of global recognition has limited formal improvement such as optimized and
mechanized production or breeding improved varieties. Despite the many advantages, fonio has
low and variable grain yield (210-1969 kg/ha) and the forage yield is unknown. Research
through field trials is needed to better understand how fonio responds to production parameters
like seeding rate and fertilizer. Recent studies in West Africa show that fertilizer can improve
fonio grain and biomass yields with the optimal range of nitrogen being between 15-40 kg/ha
(Amekli, 2013; Bakare & Ochigbo, 2003; S. Dachi et al., 2017; Gigou et al., 2009; Konaté et al.,
2021). Planting strategy such as drilling and optimal planting time also increase yields (S. Dachi
et al., 2017; M. Gueye et al., 2015). However, to the author’s knowledge, there are no published
studies recording fonio growth in the U.S.
Before it can be incorporated into U.S. crop systems, management strategies must be
explored. Specifically, it is unknown how fonio will grow in Western Michigan, if it will produce
grain, or the forage potential. There are many gaps in our knowledge of fonio growth and
potential niches. This research explored several aspects of fonio production in Michigan with a
focus on seed and fertilizer rates, but also observing varietal differences in growth and forage
quality. The purpose of this research is to characterize fonio production in a new environment
while studying specific production parameters and potential forage uses. The research
objectives were to:
1. Characterize fonio growth habit, forage yield, and forage quality in a new agroecosystem
2. Determine how fertilizer rate affects lodging, total biomass, grain yield, and plant growth
3. Determine how seeding rate affects lodging, total biomass, grain yield, plant growth
characteristics, and forage yield and quality
56
To the author’s knowledge, fonio has never been grown in the USA, thus this research will
contribute to the literature on optimized crop and forage production and will characterize fonio
growth in Michigan, USA. This research is important to better understand how fonio grows in
Michigan, potential challenges, and basic management strategies.
4.3 Methods
Study Site Preparation
Field trials measuring a) seeding rate (SS Study) b) fertilizer rate (NPK study) were
conducted over two field seasons (2021 and 2022). Only in the 2022 field season, additional
trials measuring c) interaction effects between seeding rate and nitrogen and d) observations of
six different varieties were conducted. Both years, trials were planted on an organically
managed vegetable farm called Fat Blossom in Allegan, MI with a temperate climate averaging
990 mm of rain during the growing season. A single variety and single seed lot of fonio (except
for variety trials), originating from eastern Guinea, was planted both seasons with seeding rates
adjusted on a live seed basis over the two years (80% in 2021 and 75% in 2022). The 2021 field
study was planted June 30 and the 2022 field season was planted June 1. The first year was
planted late due to heavy rainfall and availability of seed. The second year was planted earlier
to ensure full grain fill before the first frost. Both trials had four replicates and were organized in
a random complete block design, blocked by field slope. Seeds were broadcast and then lightly
raked to incorporate the seed. To combat weed pressure, fields were cultivated three days
before planting and the day of planting. Manual weeding of all distinguishable weeds (mostly
broadleaves) was completed in the third and fifth week. In the 2022 field season, another major
weed, stinkgrass (Eragrostis cilianensis), was distinguishable from fonio when flowering. After
fonio booting stage, the majority of the stinkgrass seed heads were removed to help ensure
fonio grain purity.
Experiment protocol
A) Seeding rates of 4, 8, 16, and 32 (only 2021) kg/ha were used to measure the effect
or seeding rate on fonio growth parameters and yields. In West Africa, fonio is planted at a rate
of 30-40 kg/ha with records of high seeding density of 70 kg/ha and lower rates for sowing in
rows at 18 kg/ha (Cruz et al., 2016). In this study, we tested the lower range of seeding rates.
The highest seeding rate was not included in the second season due to limited seed supply.
The plots were 1.0 x 3.0 m with 30 cm in between plots. No fertilizer or other inputs were used.
57
B) Nitrogen rates of 0, 19.5, 27.8, or 52.2 kg N/ha in one, two, or three split applications
were used to measure the effect on fonio growth parameters and yields. We incorporated with
total amounts based on previous research using a range of 0-60 N kg/ha (Amekli, 2013; Bakare
& Ochigbo, 2003; S. Dachi et al., 2017; Gigou et al., 2009). Fertilizer was applied at three rates
during planting (0, low, high) and once or twice in a split application during the growing season
(+1, +2) for a total of 7 treatments. Organic poultry manure litter was used and contained 2-3-4
NPK. The fertilized plots were 1.0 x 1.0 m and planted at a seeding rate of 24 kg/ha.
C) In 2022, a two factor study plot was planted with a low fertilizer rate (19.5 kg N/ha)
incorporated at planting and a low seeding rate (4 kg/ha). From this, a two factor comparison
was able to explore interactions between fertilizer rate and seeding rate.
D) In 2022, a variety observation was planted with five different varieties from Mali.
Varieties 1, 2, and 4 were planted at rate of 49 kg/ha and varieties 3 and 5 were planted at a
rate of 59 kg/ha. No inputs were used.
Data Collection
Prior to planting, soil at depth of 20 cm was collected in 10 random locations throughout
each block and homogenized. The samples were dried and analyzed at A&L Labs for nutrient
content. Soil texture analysis was completed by the micropipette method (adapted from Benbi et
al., 1996) in the lab.
In both years, forage quality and yield samples were taken at late booting stage (cut 1)
of development 50 days (2021) or 56 days (2022) after planting in the seeding rate trial plots
and in the variety observation plots (only 2022). In both years, another forage quality and yield
sample were taken at grain maturity 80 days (2021) or 120 days (2022) after planting (cut 2).
Only in 2022, the first forage sample quadrant was recut for forage quality and yield analysis
(cut 3). To collect all samples, a 0.5m2 quadrat was randomly thrown into a plot. All non-fonio
plants were removed and the number counted and recorded. The percentage of fonio plants at
the flowering stage was also recorded. All fonio plants were cut a few inches above soil line
(approximately 4 cm.), fresh weight was measured, and samples were dried for 72h until
consistent mass before analysis with Near Infrared Spectroscopy (NIR) which measured
micronutrients, crude protein, and fiber contents. Pre-established equations for mixed grasses
were used.
To determine plant characteristics and possible changes in plant partitioning, individual
plants sample were collected. At harvest, five (seeding rate plots) or three (nitrogen rate plots)
flowering fonio plants were randomly selected and measured for height, fresh weight, number of
58
tillers, panicles, and racemes. Each plant was cut at soil line, dried for five days, and weighed.
Total plant density was back calculated by dividing the total plot biomass by the average
individual plant. Each plot was given a lodging score (1-10) based on the extent and severity of
lodging (Bitarafan et al., 2019). For example, a lodging score of 5 would mean either the entire
plot was slightly fallen over, or one small portion of the plot was completely lodged. The entire
plot was harvested by cutting each plant at the soil line and transported to the drying location.
To dry, biomass was spread out on a tarp and turned weekly. After the biomass dried (about
four weeks), the fonio was threshed by stomping on the biomass and shaken by hand. The
grain was collected on a tarp. The dry straw was discarded, and the grain was sieved for final
yield measurements. Grain was left with hull and without complete purity from weed seed.
Statistical Analysis
The statistical software, R (Version 1.4.1717), was used to visualize and analyze the
data. Residuals were assessed for normality via data visualization. Homogeneity of variance
was examined by Levene's test with the squared and absolute values of residuals (P ≤ .05). No
transformations were needed. To measure the effect of treatment and year on the response
variables, liner mixed models were used with treatment and year as fixed effects and block as
random effects. The response variables from the experiments include whole plot measures:
above ground biomass, forage quality (SS only), lodging score, grain yield (SS only), and
individual plant parameters: plant height, plant dry mass, number of tillers, panicles, and
racemes. Fixed factors were treatment (either seeding rate or nitrogen rate), year, cut (forage
only), and the interactions. Models with and without interaction effects were compared using
AIC values and model with the lowest AIC value were used for final analysis. Analysis of
variance was used to test for significance and Tukey test was used post hoc to compare
treatments.
4.4 Results
Study Sites
Weather conditions differed between years with the recorded average temperature and
precipitation summarized in Table 4.1. Yearly data was collected from the enviroweather
platform from MSU and yearly averages from weatherspark.com. The first field season was
slightly warmer and drier than the second year, but the major difference between years was
monthly precipitation. In 2021, there was high precipitation right before planting then lower than
average for the rest of the growing season. Most notably, there was low rainfall after planting
59
and a 4 week drought from the end of August to the end of September during crucial grain-fill
period (Figure 4.1). In 2022, precipitation was close to monthly average, with a temporal drought
in the beginning of the growing season for two weeks.
Table 4.1: Total monthly precipitation and mean low-high temperature for Allegan,
Michigan during 2021 and 2022 field seasons and normal yearly averages.
Temperature (low-high oC) Total Monthly Precipitation (mm)
Year
June July August May June July August September October
20211 17-27 17-29 33.8 220 50.3 57.7 34.5 129.5
20221 13-27 16-28 16-27 115 65.3 108 89.9 34.6 62.0
Avg2 15-26 17-28 16-26 78.6 76.2 70.9 77.6 81.1 75.4
1
Yearly data from https://enviroweather.msu.edu/weathermodels/weathersummary
2
Averages from https://weatherspark.com/y/146616/Average-Weather-at-Gerald-R.-Ford-International-
Airport-Michigan-United-States-Year-Round#Figures-Temperature
300 50
250
Cumulative Rainfall
40
Daily Rainfall (mm)
200
30
150
20
(mm)
100
50 10
0 0
6/30/2021 7/31/2021 8/31/2021 9/30/2021
50
300
Cumulative Rainfall (mm)
45
40
Daily Rainfall (mm)
250
35
200 30
150 25
20
100 15
10
50
5
0 0
6/1/2022 7/1/2022 8/1/2022 9/1/2022
Figure 4.1: Daily and cumulative rainfall from planting to harvest at Fat Blossom Farm,
Allegan, MI in 2021 and 2022. Cumulative rainfall from planting to harvest in gray on the left
axis and daily rainfall in orange (2022) or blue (2021) on the right axis.
The soil texture was a major difference between study sites. The field in 2021 had higher
sand content compared to the 2022 field (Table 4.2). Between the sites for SS study, 2021 had
60
higher sand and organic matter content. Between the site for the NPK study, sand content was
much higher in the 2021 site and organic matter was lower. Between the study sites in 2021, the
NPK site had higher levels of sand (84.0% vs. 74.2%) and lower amounts of organic matter
(OM) (1.25% vs 2.05%) compared to the SS site.
Table 4.2: Soil nutrients analysis for each study and block at Fat Blossom Farm,
Allegan, MI in 2021 and 2022
K Mg Ca
Clay Sand Silt % P K Mg Ca
Year Study Block pH CEC Sat Sat Sat
% % % OM ppm ppm ppm ppm
% % %
1 4.4 73.2 22.4 5.3 2.1 109 54 90 655 6.6 2.1 11.4 49.9
2 5.2 73.6 21.1 5.3 2.0 101 36 89 511 5.8 1.6 12.8 44.1
SS
3 4.9 74.8 20.4 5.4 2.1 102 41 97 538 4.8 2.2 16.8 56
4 4.8 75.1 20.1 5.2 2.0 99 44 96 543 6.0 1.9 13.3 45
2021
1 4.0 83.9 12.1 5.6 1.1 141 57 82 556 4.8 3 14.2 57.8
2 3.2 84.2 12.5 5.7 1.2 157 48 88 464 4.4 2.8 16.8 53
NPK
3 3.6 83.8 12.6 5.6 1.3 143 48 89 660 5.4 2.3 13.8 61.5
4 3.6 84.2 12.2 5.8 1.4 166 39 80 667 5.3 1.9 12.6 62.9
* * * * * * * *
1 4.2 68.7 27.2 5.7 1.7 81 53 97 582 5.1 2.7 16 57.6
2 4.4 70.7 25.0 5.7 1.5 76 53 90 536 4.8 2.9 15.7 56.2
SS
3 5.3 65.8 28.9 5.7 1.5 81 66 105 760 6.0 2.8 14.5 62.9
4 5.7 70.7 23.7 5.6 1.4 85 57 89 492 4.5 3.2 16.3 54.1
2022
1 5.40 70.0 24.6 5.5 1.6 81 82 96 885 7.8 2.7 10.2 56.5
2 5.47 69.7 24.9 5.6 1.4 76 65 94 524 4.8 3.5 16.4 54.9
NPK
3 5.51 71.4 23.1 5.6 1.5 79 70 86 508 4.6 3.9 15.5 54.8
4 6.04 69.9 24.1 5.5 1.5 88 61 88 731 6.9 2.3 10.6 52.6
* *
* Significant differences between study sites are indicated by an asterisk at the bottom of the column
within the year.
Observation and Development
A single variety of fonio was planted for the seeding rate and fertilizer trials. Little was
known about this variety except that it was gifted by a farmer near KanKan, Guinea. From the
two field seasons, it can be determined that this variety flowers between 52-60 days and
matures 17-41 days after flowering (Table 4.3). This variation is possibly due to differences in
water availability throughout the field. Both flowering and maturation occur heterogeneously
throughout the plots and fields with flowering still occasionally occurring at harvest when most of
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the grain is mature. After grain matures, this variety also has a tendency to shatter. Lodging was
mostly observed after animals use the fields for bedding or after storms.
There was also a challenge with the variety observations and the 2021 NPK trials not
flowering. The 2021 field trials for the SS and the NPK study were in the same field but the SS
flowered and the NPK study did not. The difference between the sites of the two studies in the
same field was soil texture and slope of the field. The general observations of the single variety
are summarized in table 4.4 across all of the years and trials.
Table 4.3: Days from planting to: emergence, boot, flower, grain maturity, and
harvest for the seeding rates, fertilizer rates, and variety trials at Fat Blossom Farm,
Allegan, MI in 2021 and 2022
Planting Grain
Study Emergence Boot Flower Harvest
date Maturity
June 30 SS 8 52 60 77-81 77-81
2021 NPK 8 na na na 122
SS 6 57 57 100-121 100-121
June 1
NPK 6 50 52-60 92 92
2022
Var 7 na na na na
Table 4.4: Varietal characteristics of the single variety from Kankan, Guinea grown
at Fat Blossom Farm, Allegan, MI in 2021 and 2022. The range indicating the average
values between the two studies.
Trait
Days to Flower 50-60
Days to Grain Maturity 77-120
Avg. Lodging Score* 5-6
Avg. Plant Height (cm) 66-74
# tillers 3-4
# panicles 6-10
# racemes 30-50
Growth Habit Erect
Grain Color Brown/Tan
* Lodging score based on a 1-10 scale between complete lodging (10) and no lodging (1).
Seeding Rate Study (SS)
For above ground biomass, there was an interactive effect between year and treatment,
and year was a significant additive effect (f=9.31, p= 0.006). Between years, the total
aboveground biomass (DM) was greater in the second year (Table 4.5, Figure 4.2). In the first
year, seeding rate treatments did not affect total biomass (f=0.66, p=0.592). In the second year,
62
the highest seeding rate had a higher biomass compared to the lowest seeding rate (p=0.016).
These results may be explained by total plant density in each plot which followed a similar trend
to biomass results. For grain yield, there was no interaction between year and treatment, and
both seeding rate (f=5.48, p=0.007) and year (f=8.46, p=0.009) had additive effects. The second
year had overall higher grain yields compared to the first year. In both years, higher seeding
rates had lower grain yield. Rates of 4 and 8 kg/ha had higher grain yields compared to 16 and
32 kg/ha in 2021 and the rate of 8kg/ha had higher grain yield compared to 16 kg/ha in 2022.
Lodging was not affected by year (0.712, p= 0.410), treatment (f= 0.422, p= 0.739), or
an interaction effect in either year. Increased lodging was attributed to deer using the field to
bed according to observations made by the farmer and the lodging patterns (Figure 4.3) in the
first year and storm damage in the second year. Average plant height at grain maturity was not
different between years. In both years, the 16kg/ha seeding rate had shorter plants compared to
the lower seeding rates. In 2021, the highest seeding rate, 32 kg/ha, also had taller plants
compared to the 16kg/ha seeding rate.
Individual plant morphological traits were different between years (Table 4.5). The
number of tillers per plant were not affected by seeding rate but were significantly affected by
year (f=24.9, p<0.001) with number of tillers being higher in the first year compared to the
second year. Similarly, the number of panicles (f=16.6, p<0.001), racemes (f=30.4, p<0.001),
and individual plant dry mass (f= 18.4, p<0.001) were unaffected by treatment and higher in the
first year compared to the second year.
Table 4.5: Plot and individual plant results for seeding rate study at Fat Blossom
Farm, Allegan, MI in 2021 and 2022
Seeding Plot Data Individual Plant Data
Year Rate Lodging Height DM pani race
DM (g/m2) Grain (g/m2) tillers
(kg/ha) Score** (cm) (g) cles mes
2021 4 531 ± 33 a 5.0 14.4 ± 2.1 a 76.75 a 2.82 5.3 10.4 58.5
8 573 ± 17 a 6.0 12.1 ± 3.5 a 78.91 a 3.85 5.6 14.7 80.3
16 532 ± 42 a 5.5 5.03 ± 0.46 b 68.80 b 2.67 5.3 11.9 60.5
32 596 ± 38 a 5.5 5.81 ± 1.3 b 77.10 a 3.05 5.1 12.9 65.6
2022 4 576.5 ± 46 a 4.25 19.98 ± 2.9 ab 76.05 a 1.85 3.0 7.5 33.9
8 694.5 ± 62 ab 4.25 23.59 ± 4.6 a 74.30 a 1.60 3.2 6.6 29.1
16 893 ± 107 b 6.25 9.79 ± 1.3 b 64.00 b 1.29 3.3 6.9 25.5
* * * * * *
*Letters indicating statistically significant differences between treatments within a single year (p<0.05), no
letters indicate no statistically significant differences between treatments. Asterisks indicate differences
between years.
**Lodging score based on a 1-10 scale between complete lodging (10) and no lodging (1).
63
Figure 4.2: Total aboveground biomass yield (DM) and grain yield for seeding rates at Fat
Blossom Farm, Allegan, MI in 2021 and 2022.
64
Figure 4.3: Images of lodging at Fat Blossom Farm, Allegan, MI in 2021 field season,
comparing block 1 (lodging score of 3, left) and block 3 (lodging score of 6, right). The area
where a deer used the field for bedding is boxed in green.
Fertilizer Study (NPK)
Total above ground biomass was affected by treatment (f=3.24, p=0.0099) and year
(f=4.8545, p=0.0327) without interaction between the two terms. Between the two years, at
every treatment level, the 2022 field season had higher biomass yields. In both years, every
treatment except for the low treatment, was different from the control, meaning that additions of
fertilizer greater than 19.5 kg N/ha produced greater biomass (Figure 4.4). In 2021, compared to
the control, the highest fertilizer treatment (high+2) increased biomass by 63% and the lowest
fertilizer amount increased biomass by 23% (Figure 4.5). These results were not caused by
increased plant densities since density was the same in each plot (f=0.785 , p=0.586). Grain
yield was not collected in 2021 due to lack of flowering in the plots. In 2022, fertilizer treatment
had no effect on final grain yield (f=0.3866, p=0.878). No other factors (dry weight, plant height,
etc.) were correlated to final grain yield.
Lodging was not affected by treatment or year. Plant height was affected by year (f=113,
p<0.0001) and not affected by treatment (f=0.93, p=0.48) or an interaction between terms. Plant
height was greater at every treatment level in the 2022 field season.
Individual plant morphological traits were not different between years (Table 4.6). The
number of tillers per plant were not affected by fertilizer rate (f=0.703, p=0.65) or year (f=3.39,
65
p=0.072). Similarly, the number of panicles (f=0.733, p=0.29), racemes (f=0.783, p=0.593), and
individual plant dry mass (f=0.593, p=0.733) were unaffected by treatment and year.
Table 4.6: Plot and plant results from the fertilizer trials at Fat Blossom Farm,
Allegan, MI in 2021 and 2022
Plot Data Individual Plant Data
Fertilizer DM Grain (g/m2)
Year Heigh DM till pani race
Rate 2
(g/m ) ± SE ± SE t (cm) (g) ers cles mes
0 572.7 ± 43.9a 44.8 1.43 4.6 9.5 38
Low 704.2 ± 30.5 ab 43.1 1.08 2.9 4.8 22
Low+1 872.0 ± 38.2b 51.0 1.31 3.6 5.3 26
2021 Low+2 865.2 ± 61.9b 49.0 1.09 4.1 7.5 34
High 848.2 ± 27.7b 43.7 0.81 4.3 5.7 30
High+1 935.5 ± 52.4b 49.9 0.93 3.0 5.3 21
High+2 937.8 ± 61.1b 56.0 2.18 5.1 11.8 60
2022 0 732.5 ± 49.5a 23.27 ± 5.5 a 73.1 1.26 2.9 5.1 25.4
Low 920.0 ± 40.8ab 21.08 ± 3.5 a 81.8 1.59 3.5 4.4 24.0
Low+1 903.8 ± 59.7b 23.58 ± 5.9 a 79.8 1.43 2.9 5.0 26.7
Low+2 943.8 ± 46.2b 22.31 ± 3.8 a 82.0 1.98 3.8 4.9 26.6
High 1028 ± 21.7b 17.24 ± 1.6 a 88.4 1.85 3.0 4.3 24.1
High+1 973.8 ± 100b 21.66 ± 3.9 a 86.3 2.21 3.3 5.3 29.3
High+2 921.3 ± 30.7b 23.28 ± 1.8 a 88.3 1.92 2.8 4.8 26.7
* *
* Average dry mass (DM) and distribution and individual plant parameters at each fertilizer rate.
Letters indicate differences in treatments within a field season.
66
=
Figure 4.4: Total aboveground biomass yield for each fertilizer treatment at Fat Blossom
Farm, Allegan, MI in 2021 and 2022
0 Low Low+2 Low+1 High High+1 High+2
37 days
52 days
94 days
Figure 4.5: A single replicate in 2021 season over three time points: at +1 and +2
fertilization and at harvest at Fat Blossom Farm, Allegan, Mi
67
Interaction between seeding rate and fertilizer rate
In the exploratory study c) investigating the interaction between seeding rate and
fertilizer, we had one year of data from 2022. For total above ground biomass yield, there was
no interactive effect between seeding rate and nitrogen rate. Seeding rate, regardless of
fertilizer rate, increased total aboveground biomass (Table 4.7). Fertilizer only increased
biomass at higher seeding rates. Grain yield, lodging, and individual plant traits were all
unaffected by fertilizer or seeding rate.
Table 4.7: Plot and plant results from the fertilizer x seeding rate trials at Fat
Blossom Farm, Allegan, MI in 2022
Grain Ind.
Fertilizer Seeding Dry Mass Lodge #till #pan # rac
(g/m2) Height Dry
rate Rate (g/m2) Score** ers icles emes
mass
0 4 576.50±46.0 a 19.98±2.9 4.25 76.05 2.95 7.50 33.90 1.85
0 24 732.50±42.9 b 23.27±4.8 6.25 73.08 2.92 5.08 25.42 1.26
Low 4 696.25±31.8 ab 19.92±5.2 6.50 81.50 4.50 6.75 39.33 2.11
Low 24 920.00±35.4 c 21.08±3.0 6.75 81.75 3.50 4.42 24.00 1.59
*Letters indicate differences in treatments within a field season. No letters demonstrate no difference
between treatments.
**Lodging score based on a 1-10 scale between complete lodging (10) and no lodging (1).
Forage Yield and Quality
In both years, seeding rate had no effect on forage yields (f=1.827, p=0.155), crude
protein (f=1.94, p=0.155), fiber content, or forage quality (f=2.32, p=0.110), and thus, seeding
rate effect was excluded from the analysis. Forage yield (DM) was affected by cut (f=122,
p<0.001), year (f=8.78, p=0.0047), and the interaction (f=18.36, p<0.001). Forage yield was
greater during the 2021 field season compared to the 2022 field season (Table 4.8). Within each
year, biomass increased with each cut. The recut of the first sample (cut 2) had lower biomass
compared to an uncut sample (cut 3). The yield for the first cut on average across seeding rates
was 123 g/m2 and 45.8 g/m2 in the first and second year (Figure 4.6). For crude protein, cut
(f=138, p<0.001) and year (f=151, p<0.001) both had an affect with no interaction between
terms. At each cut, crude protein was higher in 2021. For both years, the first cut had the
highest crude protein. Within the second year, there was no difference in the crude protein
content of the second cut and the recut of the first cut. For fiber content (both NDF and ADF),
68
cut (f=453, p<0.001), year (f=130, p<0.001), and the interaction (f=360, p<0.001) had an effect.
Fiber content was greater in the second year and increased with each cut. Similarly, relative
forage quality was affected by cut (f=100, p<0.001), year (f=36.6, p<0.001), and the interaction
(f=72.1, p<0.001). Between years, the second year had higher forage quality at the first cut (151
vs. 134), but the first year had higher forage quality at the second cut. In both years, the forage
quality decreased with subsequent cuttings. In the second year, the recut had higher forage
quality compared to the uncut sample.
Table 4.8: Forage Potential Results from the seeding rate study at Fat Blossom
Farm, Allegan, MI in 2021 and 2022
Year Cut #** DM (g/m2) CP ADF NDF RFQ
1 123 ± 7.8 a 15.3 ± 0.22 a 31.91 ± 0.14 a 60.0± 0.13 a 134 ± 0.42 a
2021
2 558 ± 19 b 9.65 ± 0.39 b 35.6 ± 0.27 b 62.7 ± 0.41 b 121 ± 2.0 b
1 45.8 ± 2.1 a 11.0 ± 0.17 a 29.7 ± 0.31 a 53.7 ± 0.49 a 151 ± 2.1a
2022 2 288 ± 26 c 4.40 ± 0.27 b 41.6 ± 0.24 c 70.7 ± 0.27 c 107 ± 1.8 c
3 225 ± 24 b 5.05 ± 0.35 b 40.5 ± 0.17 b 68.8 ± 0.36 b 118 ± 2.5 b
* * * * *
*Dry Mass (g/m2) and NIRS forage quality results including: % Crude Protein (CP), % Acid Detergent
Fiber (ADF), % Neutral Detergent Fiber (NDF), and Relative Forage Quality (RFQ). ± the standard error
(SE). Letters indicate differences within years between cuts, asterisks indicate differences between years.
**Cut 1 was taken at the booting stage for optimal forage quality, Cut 2 was a separate cutting taken at
grain harvest for optimal forage yield, Cut 3 was a re-cutting of Cut 1 taken at grain harvest
69
mixed grass/legume
other legume
alfalfa
sorghum
small grain
fonio 2022
fonio 2021
0 5 10 15 20 25
% CP
mixed grass/legume
other legume
alfalfa
sorghum
small grain
fonio 2022
fonio 2021
0 50 100 150 200
RFQ
Crabgrass
Teff
Red Clover
Perennial Grass
alfalfa
fonio 2022
fonio 2021
0 5000 10000 15000 20000
DM (kg/ha)
Figure 4.6: Comparing crude protein (CP)1, relative forage quality (RFQ) 1, and dry matter
(DM, kg/ha) 2 across forage types. The average (end of bar) and typical expected range (black
bars).
70
1
RFQ and CP data from various forage species submitted to the UGA Feed and Environmental Water
Laboratory from July 2003 to February 2011. Data is adapted from Understanding and Improving Forage
Quality | UGA Cooperative Extension. Data from our field trials added with calculated average values of
fonio RFQ and CP and minimum and maximum indicated by the bars in 2021 field season and 2022 field
season.
2
DM data is adapted from the MSU forage connect variety trial results in Michigan during the years of
2019-2021. Data from our field trials added with calculated average values of fonio DM and minimum and
maximum indicated by the bars in 2021 field season and 2022 field season .
Variety Trials
Five fonio varieties from Mali were planted for observation and tested for forage quality
without replication in the second field season (Table 4.9). No flowering was observed in any of
the plots for unknown reasons. Variety #4 suffered from an unknown leaf curling disease that
plant diagnostics could not identify. Lodging was also observed in all varieties but not scored.
The varieties had similar forage quality and biomass yield with variety 5 appearing to have
higher biomass yield and quality in the first cut and higher quality but lower biomass yield in the
second cut (not a re-cut). Generally, there were differences observed in forage quality and yield
between varieties.
Table 4.9: Forage Potential Results of different varieties from observational trial
with single replicate in 2022
DM
Cut Variety CP ADF NDF RFQ
(g/m2)
1 58.29 11.02 29.04 51.00 142.3
2 63.37 8.74 30.49 53.53 143.9
1 3 54.87 9.06 29.62 53.16 147.5
4 76.25 9.70 29.27 52.79 147.5
5 79.56 9.85 29.25 53.09 148.2
1 497.5 2.22 39.50 70.63 99.6
2 396.1 1.52 39.56 71.99 93.5
2 3 355.5 1.77 40.54 72.59 92.4
4 437.4 3.70 40.38 69.67 109.4
5 286.4 4.53 41.78 70.35 112.3
*Dry Mass (g/m2) and NIRS forage quality results including: % Crude Protein (CP), % Acid Detergent
Fiber (ADF), % Neutral Detergent Fiber (NDF), and Relative Forage Quality (RFQ). Varieties 1, 2, and 4
were planted at rate of 49 kg/ha and varieties 3 and 5 were planted at a rate of 59 kg/ha. Cut 1 was taken
at booting stage. Cut 2 was a separate cutting taken before frost.
71
4.5 Discussion
Seeding Rate Study
We examined the effect of seeding rate on plant growth and yields to determine optimal
management practices in Michigan. According to Cruz et al (2016), fonio is traditionally planted
at a rate of 30-40 kg/ha with records of high seeding density of 70 kg/ha and lower rates for
sowing in rows at 18 kg/ha. In this study, we tested the lower range of seeding rates. The
seeding rate increased above ground biomass only in the second year which may be explained
by low precipitation in the first field season limiting plant growth. Other small grains also
demonstrate increased above ground biomass as seeding rate increased (Baker, 1982; El-
Lattief, 2014) most likely due to overall greater plant density per plot. Contrary to fonio studies
comparing biomass and grain yields (Amekli, 2013) and other small grains (Baker, 1982; Dorval
et al., 2015; El-Lattief, 2014; Schillinger, 2005), fonio grain yield was reduced by seeding rate
and negatively correlated to biomass. Only tef grain and biomass yields have been observed to
increase at lower sowing rates (Arefaine et al., 2020). The grain results are also not explained
by plant morphological traits measured. In the second field season, increased reproductive
panicles and racemes at lower plant densities were observed but were not statistically
significant or observed in the first year. On average, the number of reproductive tillers, panicles,
and racemes were greater in the first field season, but did not correspond to greater grain
yields.
We hypothesized that both grain and biomass yields would increase as seeding rate
increased but we found that only biomass was improved with greater seeding rates. These
results were not explained by morphological characteristics but may be due to overall low grain
yields and a different climate from the variety’s origin. The seed variety grown originated from
Kankan, Guinea which averages 590 mm of precipitation during the growing season. The
average precipitation in Allegan, MI is 306 mm, almost half and was exasperated by temporal
droughts throughout the growing season especially in 2021. This possibly led to overall low
yields especially compared to average yields of fonio in Guinea (Figure 1.1). These results also
may be affected by the different planting dates between the two years. A later planting date
allowed for longer amount of time for total growth and biomass accumulation in the first season.
As hypothesized, average individual plant height was also impacted by seeding rate, but
lodging was not. The seeding rate of 16 kg/ha produced shorter plants compared to the other
seeding rates but did not correlate to less lodging. This is contrary to many findings where
lodging and plant height are positively correlated (Arefaine et al., 2020; Dorval et al., 2015). The
72
greater factors affecting lodging in the fonio plots were storm winds and animals. To further
understand fonio response to lodging and height, more controlled studies may need to be done.
Nitrogen Rate Study
We examined the effect of fertilizer rate on plant growth and yields to determine optimal
management practices in Michigan. In both field seasons, a positive trend was observed
between fertilization and biomass. Fertilized plots with nitrogen greater than 19.5 kg N/ha had
statistically significant different above ground biomass compared with those with less (0 and low
treatments). These results are similar to studies from West Africa. Gigou et al. (2009) found that
fonio responded to inorganic NPK fertilizers at low levels of fertilization (30 kg/ha) which
increased grain yields by 22%. Amekli (2013) also determined that increased grain yields
directly correlated with an increase of biomass. Grain yields from our study were not affected by
fertilization or correlated with biomass yields. The lack of statistical differences between
treatments in grain yield may be explained by the precipitation challenges explained above and
demonstrates challenges with growing crops in new agroecosystems.
The results demonstrate an important risk when fertilizing fonio. The regions where fonio
is grown, especially for food security and sustenance, are prone to drought. Fertilizer may have
little to no affect during drought and there may not be a corresponding increase in grain yield.
Using fertilizer does not guarantee higher yields since it is dependent on rainfall and many other
factors. Maji & Imolehim (2013) also determined fertilization without fungicide application
increased the severity of brown spot disease and decreased grain yields. Fertilizer may not
guarantee increased grain yield for sustenance farmers because of nutrient availability also
depending on soil moisture and precipitation.
Challenges Across Studies
Flowering is a challenge when growing a crop in a new environment because there are
many biological and environmental factors that control grain development. In 2021, the fertilized
field trial did not flower, but the seeding rate study, in the same field with the same climate and
precipitation, did successfully flower and produce grain. The major differences between the two
trials were soil organic matter and texture, two factors heavily influencing water holding
capacity. The slope of the field may have drained water into the seeding rate plots. Low rainfall
during the beginning of the season made water a limiting factor during crucial growth stages.
We hypothesize that the nitrogen plots did not flower in 2021 due to lack of water, also possibly
limiting the availability of nutrients. The following season, there was much more rainfall
73
throughout the season (Fig. 4.1) and the plots did not experience a lack of flowering. Similar
conclusions were found by Gigou et al. (2009) at a field site with similar rainfall and soil
parameters that fertilizer had less effect on total above ground biomass when water was
limiting. In the 2022 field season, the fertilized plots flowered and produced grain, but an
increase of biomass did not correlate to an increase of grain yield.
Grain yields may be impacted by the challenge of harvesting and processing fonio
grains. Overall grain yields (0-260 kg/ha) in both seasons were extremely low especially
compared to yields in West Africa (275-1175 kg/ha) (Figure 1.1). Fonio is known to shatter at
grain maturity (Bakare, 2005; Cruz et al., 2016). During both field seasons, grain matured at
different rates, so a plot was harvested when the majority was at grain maturity. However, there
is a possibility that grain was lost due to shattering and plots were harvested outside of peak
grain maturity. There were also challenges with separating fonio grains from other weed seeds.
The size of fonio grains is similar to amaranth and many other common weeds meaning that
fonio grain was challenging to separate by sieving or by weight. This may lead to inflated fonio
grain yield measurements and possibly greater grain yields in plots with more weeds. Future
studies may prevent these challenges by more tedious removal of weeds from the plots or an
effort to harvest fonio seed heads before cutting for biomass yield. Learning from West African
producers may have also aided in the process but was not feasible for this study. There may
also be opportunities to learn and adapt mechanized processes from U.S. farmers who process
comparable seed.
Forage Potential
The potential of fonio as a forage crop in the United States and elsewhere, depends on
the quality of the stover and the biomass yield. In this study, we assessed forage quality at
different cuts in the seeding rate study and measured the dry biomass. During the first season,
at optimal forage quality, fonio forage had higher quality and crude protein compared to small
grains, annual ryegrass, orchard grass, Bahia grass, Bermuda grass, pearl millet, sorghum, and
peanut vines (UGA Cooperative Extension). Forage quality was only lower compared to
legumes and alfalfa, but yields were low at 1236 kg/ha (0.5 tons/acre). In the second field
season, quality was comparable to alfalfa forage quality with an RFQ of 151, but much lower
yields 470 kg/ha (0.19 tons/acre). According to the USDA, forage yields for annual summer
grasses such as millets and crabgrass are on average between 4-8 tons/acre. Forage yields of
fonio in both years were well below the average for summer grasses. However, compared to
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other annuals grown in Michigan, like tef grass and crabgrass, the forage yield was comparable
(Fig. 4.6).
We also assessed forage yield and quality at grain harvest, the highest forage yield
potential. In the first year, the second cut was comparable in quality to other grasses according
to UGA Cooperative Extension with RFQ of 121, and yields were higher at 5585 kg/ha (2.26
tons/acre). In the second field season, forage quality (RFQ= 118) and yield were lower-
2249kg/ha (0.91 tons/acre). The difference between years is unexpected since year 2 had
greater rainfall (Fig.1) and was planted earlier. In the second year, a recut (cut 3) was also
taken of the original first cut to test the possibility of cutting fonio twice for forage. The recut had
lower yield than the second cut but higher quality. Total yield for two cuts (1 cut at booting stage
and a recut at grain maturity) was 2718 kg/ha (1.1 ton/acre) which is low compared to other
summer grasses, but comparable to summer annuals grown in Michigan.
We hypothesized that seeding rate would increase forage yields but found that forage
yield and quality were unaffected by seeding rate. These results are similar to spring oat grown
for forage in the central great plains where seeding rate did not affect forage yield or quality
(Obour et al., 2019). Forage sorghum demonstrated that an increase of seeding rate decreased
stalk thickness and improved quality but did not having an effect on forage yield (Mekasha et al.,
2022). It may be concluded that forage yield is not affected by seeding rate, but plant
partitioning at higher levels of competition may affect forage quality. For fonio forage production,
low seeding rates are acceptable. The five varieties tested demonstrated that variety may
influence forage yield, quality, and the potential for multiple cuts. The varieties yielded 544-791
kg/ha (0.22-0.32 ton/acre) in the first cut and 2820-4890 kg/ha (1.14-1.98 ton/acre) in the
second cut. These variety trials were unreplicated over a single field season, so more robust
variety trials are needed to determine varieties with optimal qualities for forage yield and quality.
Fonio being a high quality, but low yielding forage crop has multiple implications for
farmers globally. In West Africa, fonio is used both as a forage crop and as a grain crop for
animals (C. I. Ukim et al., 2021). As a dual use crop for grain and hay or forage, fonio may have
increased economic benefit for farmers. Fonio can be grazed or cut at the booting stage for
optimal quality while still producing a grain crop. The leftover hay may also be used for animal
feed. As a grazed crop, we still do not know the regrowth potential after grazing or how fonio
may perform in a pasture mixture. In the U.S., fonio may be grown as a second crop and benefit
farmers since it has low nutrient requirement and quick maturing varieties. These are the first
forage quality assessments. Forage quality measurements are needed across varieties which
75
may inform breeding efforts to improve the forage yield. Yield will need improvement to better
compare with summer annuals if fonio is to be grown globally as a forage crop.
Management Recommendations
Based on these studies there may be different management strategies based on the
intended end use for biomass or grain yields. There were no interactions between a low and
high seeding rate and a zero or low fertilizer rate. The only fonio yield affected by fertilizer and
seeding rate was above ground biomass, which increased by seeding rate, but only increased
by fertilizer at higher planting densities, indicating that farmers may benefit less from fertilizer
when fonio is planted at low rates. There was also no corresponding increase in grain yield
which is contrary to the correlation between fonio biomass and grain yield established by Amekli
(2013) but may be impacted by the challenges previously stated in collecting grain yields. More
field seasons are needed to verify these results, but these preliminary findings support that
farmers may not benefit from fertilizing fonio sown at low rates and sowing fertilizer at a high
rate may increase above ground biomass but may not increase grain yield. We recommend that
when growing fonio for grain, low seeding rates be used with no fertilization and when growing
fonio for forage, high seeding rates be used with low amounts of fertilization.
4.6 Conclusion
The purpose of this research was to characterize several fonio varieties’ growth and
forage potential in Michigan and begin studying the effect of fertilizer and seeding rates on
yields. There are many challenges associated with growing fonio in a new environment
including: lack of flowering, weed pressure, and grain harvest and purity. Our findings suggest
that fonio may be planted at low seeding rates with zero or low fertilizer inputs. Higher seeding
rates may benefit from low amounts of fertilizer especially for better aboveground biomass. As a
forage, fonio is high in quality and may be cut several times during the growing season but has
much lower biomass yield compared to other annual grasses. Preliminary variety trials indicate
that forage quality, biomass yield, and regrowth ability may be affected by variety. Further
research on fonio is needed to better understand its potential as a cover crop or forage mixture
in U.S. agricultural systems.
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5. CONCLUSIONS
Increased biodiversity greatly benefits agroecosystems by improving function and
stability over time. Management for resiliency is especially needed as climate change puts more
strain on production. Minor crops offer a resource to fill niches where major crops cannot and an
opportunity to increase diversity. Fonio may be one of these crops to contribute to global
diversity and may fill a niche as a drought tolerant, adaptable, nutritious small grain for human
and animal consumption. There is little known about fonio production beyond the West African
context and research on fonio had not yet been synthesized to find major gaps in knowledge.
The interdisciplinary scoping literature review revealed areas with little research and how
that research may benefit other disciplines, leading to insights for innovation. There are major
gaps in our knowledge of production, specifically in breeding and improved production practices.
While several management practices have been tested, such as planting date, fertilization, and
sowing method, these studies have not yet been tested across varieties or environments.
Mechanization of processing may also benefit producers by reducing the labor needed to
prepare the grain for consumption. Breeding programs will also benefit producers by reducing
the amount of shattering and grain loss which occurs during harvest. There is abundant
opportunity to increase the market for fonio products especially in urban areas, but first,
production must increase to support the growing demand.
To test fonio production in a new environment, the field study documented growth of
fonio in Western Michigan, laying the foundation for growing fonio in the Midwest and identifying
how specific management practices may affect fonio growth and yields. The study found that
low seeding rates and low fertilization improved total biomass yields and had little to no effect on
grain yield. Relative forage quality was high (130-150) when cut for optimal forage quality, but
yields were low (2249-5585 kg/ha) compared to alfalfa and clover. Fonio may be cut several
times for forage, but quality is reduced with subsequent cuttings. From these trials, it can be
concluded that fonio may be grown with relatively few inputs but would benefit from breeding to
improve forage yields. The trials also emphasized several challenges with growing fonio in
Michigan. Growing fonio as a grain crop may be challenging due to inconsistent flowering and
grain processing challenges. More research will also be needed to test the resistance of fonio to
drought since this is widely accepted, but not necessarily confirmed in our trials. Further
research may test specific niches for fonio in U.S. cropping systems.
The next steps in fonio research must be to support increased production and breeding
programs with a specific focus on improving fonio for West African producers. A breeding
77
program to help reduce grain loss such as shattering, and lodging is most likely the first step in
varietal improvement. Increasing grain size may also improve yields but may affect other factors
such as grain quality and storability. New varieties must also be tested across many
agroecosystems, even globally. More research could benefit U.S. cropping systems by finding
niches for fonio and increasing total number of crops grown per region. As research continues,
every effort must be made to empower fonio producers and support the cultural value of fonio.
In breeding, this looks like continued evaluation of farmer and consumer needs. Greater
documentation of fonio consumption is needed throughout West Africa. Participatory breeding
may also produce valuable insights from farmers and aid in adoption. Continued strengthening
of seed systems can help preserve genetic diversity of fonio and disseminate improved
varieties. When technologies are shared among farmers, programs must intentionally give
opportunity for historically disempowered farmers to participate. When marketing fonio in new
global markets or as new products, researchers, and consumers can play a part in responsible
use of genetic resources and products. Bottom-up regulation from farmers through collective
action or NGO’s may also help protect fonio producers.
78
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