Natural Sciences
Emergence of a Viral Disease Epidemic of Sweet potato in Eastern Africa is Imminent – Plant virologists warn
Published
3 years agoon

When COVID-19 emerged in 2019, it first remained temporarily localized as an epidemic in Huan China until it broke Chinese national and Asian continental borders attaining a global pandemic status. With COVID-19 and other epidemics and/or pandemics of humans and livestock, it is easier to explain the obvious impact on humans and/or livestock.

What about virus epidemics and pandemics that affect plants?
Virus epidemics and pandemics that affect plants cause significant disruptions in food production. In fact, the emergence and re-emergence of extremely dangerous plant viruses is the major pain to the 21st century agriculture and sustainable food production.
Viral diseases of plants lead to complex outcomes of food shortages, economic meltdown and loss of lives especially when the affected crop plants are central to the economics and livelihoods of the people. Think of what would happen if an epidemic caused significant effect or eliminated crops like maize, banana, cassava or sweetpotato from sub-Saharan Africa.

Most viruses affecting livestock and humans are spread and transmitted by means of physical contact between hosts. Furthermore, human and livestock hosts are mobile. On the other hand, plants do not move and must thus depend on other agents (vectors) for the spread of viruses between hosts. Consequently, over 98% of plant viruses are transmitted by vectors. These vectors may include arthropods (especially insects), nematodes, fungi, and parasitic plants. In plants, a virus disease epidemic becomes optimized by the ability of vectors facilitating fast spread of the virus in an agro-ecosystem.

New study predicts emergence of disease epidemic in sweetpotato
A new study led by Prof. Arthur Tugume at the Department of Plant Sciences, Microbiology and Biotechnology in the College of Natural Sciences at Makerere University predicts potential of emergence of a new disease epidemic in sweetpotato.
The study recognizes that by default, viruses are not constrained to perpetually occupy a single ecological niche. This is because viruses have inherent genetic and evolutionary abilities that enable their acquisition of better fitness to always expand their host and geographical ranges.
While viruses enjoy such fitness advantages in nature, the current study points out unique examples where some viruses in eastern Africa behave awkwardly. These viruses first hid under the guise of being less important and seemingly of no impact in cropping systems, and all of a sudden hell broke loose and the entire farming system was caught unaware.
The study titled; “Endemism and reemergence potential of the ipomovirus Sweetpotato mild mottle virus (family Potyviridae) in Eastern Africa: half a century of mystery” shows that a virus, named Sweetpotato mild mottle virus (SPMMV) was first detected in eastern Africa in the 1970’s. SPMMV is currently the third most prevalent virus of sweepotato in the region and is most destructive in mixed infections with other viruses. Very strangely, SPMMV is geographically restricted to eastern Africa (confirmed only in Uganda, Kenya and Tanzania). This sharply contradicts other known sweetpotato viruses that occur globally wherever sweetpotato is grown.
According to Dr. Deusdedith Mbanzibwa a co-author and Head of Disease Control Unit at Tanzanian Agricultural Research Institute, “it is a half-century-old mystery how SPMMV remains localized in East Africa while other viruses on sweetpotato show a global incidence.” Dr. Mbanzibwa adds, “it is important we generate basic information on this virus as soon as possible such that immediately an epidemic explodes we can counter-it based on the information we have as opposed to being caught unaware of the destructive effects of an epidemic.”

Why should we be concerned about outbreak of an epidemic on sweetpotato?
Plant disease epidemics cause a breakdown in food production and availability in communities that depend on the affected plants as food staples. If sweetpotato production breaks down, it directly translates into limited food supplies in local communities that depend on it for food.
According to Dr. Titus Alicai, a senior virologist and co-author based at the National Crops Resources Research Institute (NaCRRI), Namulonge Uganda, “between the 1980’s-1990’s, a devastating viral pandemic (cassava mosaic disease, CMD) destroyed cassava cropping systems in eastern Africa. As a result, dozens of lives were lost because many communities solely depended on cassava for food and income”. Although these viruses had been detected in the region as early as 1800’s, it was not until the 1980s that destructive effects of the disease became apparent.
Dr. Alicai adds that, “when cassava got destroyed by CMD, sweetpotato became an immediate alternative to millions of hungry people, but now sweetpotato is also burdened with diseases arising from viruses.” Despite heavy investments in disease control in cassava, the CMD pandemic is not completely controlled.
Why is eastern Africa important with respect to emergence of plant viruses?
According to this study, SPMMV is not the first unique virus to emerge or reemerge in eastern Africa. An earlier study by Prof. Tugume and colleagues published in 2010 in Molecular Ecology showed that East Africa is a hotspot for the evolutionary diversification of yet another virus in sweetpotato, named Sweetpotato feathery mottle virus (SPFMV).
Many studies show this region as the home of epidemics in rice caused by Rice yellow mottle virus (RYMV). Highly lethal strains of Cassava mosaic geminiviruses (CMGs) causing CMD initiated this epidemic from eastern Africa. Unique strains of Sweetpotato chlorotic stunt virus (SPCSV) are incident in eastern Africa. SPCSV is unique because it almost indiscriminately enhances the viral concentrations of other co-infecting viruses in sweetpotato. The list of damaging plant viruses or their strains that have emerged from eastern African region is long and cannot not be exhausted here.
Biologists studying diversity of plants and animals have also coded eastern Africa as a home of the “hottest hotspot of biodiversity” of endemic land plants and animals located in Eastern Arc Mountains of Kenya and Tanzania. The features of eastern African ecosystem that incite virus epidemics in plants are not fully known. However, it is suspected that the microclimatic changes as modulated by the Indian Ocean dipole could be one of the drivers of virus emergence and reemergence in the region.
Emergence of SPMMV in sweetpotato closely resemble that of viruses in cassava
The current study specifically compared SPMMV with two viruses infecting cassava: cassava brown streak virus, and Ugandan cassava brown streak virus, collectively called cassava brown streak ipomoviruses (CBSIs).
CBSIs first appeared in the 1930’s in coastal east Africa but went completely silent for about 70 years and then suddenly re-emerged in late 1990’s-mid 2000’s to what is today the single largest threat to cassava production in Africa. CBSIs cause a disease called “cassava brown streak disease” (CBSD). Between 1930’s-1990’s, the CBSIs were able to “amass ability” to cause untold suffering of cassava cropping from 2000’s to-date.
Similarly, since the 1970’s when SPMMV was first reported in the region, the virus has been “amassing an ability” to cause more disease in sweetpotato. This is implied directly by evidence of positive selection found in its genes. Positive selection refers to “survival for the fittest” in Charles Darwin’s terms to mean that, individual virus particles in the SPMMV population are becoming more and more highly efficient in causing disease. Today, half a century later, one can argue that it is about time that the SPMMV will incite a new disease.
Note that both SPMMV and CBSIs belong to the same taxonomic grouping of plant viruses in a genus called “Ipomovirus”, and family called “Potyviridae” which is the largest family of plant viruses with RNA genomes. Hence, similarities in the disease spread between SPMMV and CBSIs are not unexpected.
Expert guide to sweetpotato virology research for preparations against the epidemic
“If we can generate advance basic scientific information concerning how the SPMMV infects sweetpotato plants, how it is transmitted by vectors between plants, and circumstances fostering field spread of the virus, we will be better prepared to handle an epidemic that may arise from the virus,” Prof. Tugume states.
He adds: “we should be able to undertake these studies quickly because SPMMV is naturally incident on sweetpotatoes in Uganda, Kenya and Tanzania in levels causing less damage on its own at the moment, although this damage escalates when the virus occurs in multiple infections.”
The co-authors highlight serious research gaps for critical research investment in sweetpotato virology. Vector transmission is such an important aspect of virus disease epidemics of plants. Therefore, this study has proposed three possible explanations to account for the virus-vector and virus-host relationships of SPMMV. These are called hypothesis #1, #2, and #3, built in a fashion that allows experimentation to prove them right or wrong:
- Hypothesis #1: “SPMMV is opportunistically aphid-transmitted with potyvirus SPFMV as a helper virus.” This hypothesis postulates that SPMMV is transmitted by “chance” by a group of insects called aphids but this happens when SPMMV co-infects sweetpotatoes with another virus, SPFMV that is transmitted efficiently by aphids.
- Hypothesis #2: “Vector-mediated transmission of SPMMV is modulated by synergism with SPCSV.” Under this hypothesis, the concentrations of SPMMV in plant tissues increases when it co-infects with another virus, SPCSV, and this increases chances of SPMMV being picked by insects for transmission to other plants.
- Hypothesis #3: “SPMMV tropism and histo-localization changes upon co-infection with SPCSV.” How much viral concentration is found in different plant tissues can change depending on which tissues that are preferred by the virus. Certain plant viruses prefer to stay and multiply from certain tissues than others.
Dr. Alicai said, “If we can determine vector transmission dynamics including the actual vector(s) transmitting SPMMV, which plant tissues the virus prefers for its multiplication, and generate many complete sequences of SPMMV genomes, we will be better prepared for epidemics arising from the emergence of SPMMV”.
Sweetpotato is a vegetatively propagated plant in which the top-most 15-40cm part of the plant is used for initiating the new crop, a practice that is the main route of accelerated of virus spread in the cropping system. Also, insects especially aphids and whiteflies are the major vectors transmitting viruses in sweetpotato. The research gaps highlighted with respect to SPMMV require urgent attention for advance preparation in case an epidemic breaks loose on the sweetpotato cropping system.
This study has been published by the American Phytopathological Society (APS) in an open access journal, “Phytobiomes Journal”, and is freely accessible on https://apsjournals.apsnet.org/doi/10.1094/PBIOMES-05-22-0031-RVW
For more details, please contact;
Prof. Arthur Tugume
Lead Scientist
Department of Plant Sciences, Microbiology and Biotechnology
College of Natural Sciences (CoNAS)
Makerere University
Email: arthur.tugume@mak.ac.ug
Tel: +256772514841
Dr. Deusdedith Mbanzibwa
Tanzania Agricultural Research Institute (TARI),
Biosciences Centre, Dodoma, Tanzania
Email: d.mbanzibwa@yahoo.co.uk
Email: Tel: +255755881758
Dr. Titus Alicai
Root Crops Research Program
National Crops Resources Research Institute (NaCRRI)
National Agricultural Research Organization (NARO)
Email: talicai@hotmail.com
Tel: +256772970585
Hasifa Kabejja
Principal Communication Officer
College of Natural Sciences
Makerere University
Email: pr.cns@mak.ac.ug
Tel: +256774904211
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Natural Sciences
Enhancing Data Quality: NutriFishPlus Research Assistants Retooled Ahead of Baseline Survey
Published
2 weeks agoon
January 20, 2026
The NutriFishPlus Project team successfully conducted a one-day capacity-building training for its research assistants, centered on the Monitoring, Evaluation, and Learning (MEL) framework and the household survey tool. Held on 13th January 2026 at Ridar Hotel Seeta in Mukono District, the training aimed to improve data collection, analysis, and learning processes to support evidence-based project implementation and impact assessment.
It was officially opened by Dr. Jackson Efitre, the Principal Investigator, who introduced the research assistants to the overall goal of the NutriFishPlus Project and the guiding principles of the International Development Research Centre (IDRC) Scaling Approach. In his remarks, Dr Efitre emphasized the importance of high-quality data in tracking progress, informing adaptive management, and scaling impact.

NutriFishPlus builds on the significant achievements of the ground-breaking NutriFish Project (2019–2023), which was funded by IDRC and the Australian Centre for International Agricultural Research (ACIAR) through the Cultivate Africa’s Future Fund Phase II (CultiAF2). Officially launched on 28th October 2025, the two-year and eight-month project seeks to scale proven innovations while deepening nutrition, livelihood, and gender outcomes within small pelagic fisheries.
The project aims to:
- Scale up the adoption of improved and sustainable fish processing technologies, including solar tent driers and raised racks, to reach new fishing communities across Uganda.
- Enhance market access and strengthen supply chain linkages for high-quality fish and fish-based products.
- Empower women and other vulnerable groups while strengthening community resilience through diversified income-generating activities.

Expected Outcomes:
NutriFishPlus is expected to deliver multiple socioeconomic and environmental benefits, including:
- Improved incomes and livelihoods for marginalized fishing communities, particularly women and youth.
- Enhanced health and nutrition outcomes through the development of diversified, market-embedded fish products.
- Establishment of sustainable and replicable fish processing and marketing models across Uganda and the wider East African region.
- Increased participation of women and youth in decision-making and benefit-sharing within the Small Pelagic Fishes (SPFs) value chain.
- Improved socioeconomic conditions and ecosystem health through participatory and scalable approaches.

Project Implementation and Partnerships
The project is implemented by the Department of Zoology, Entomology, and Fisheries Sciences at Makerere University, in partnership with Nutreal Ltd and Kati Farms Ltd under a public-private partnership. Project activities will be carried out at five landing sites across Lakes Victoria, Kyoga, and Albert.

The project team comprises:
- Dr. Jackson Efitre (Principal Investigator)
- Dr. Dorothy Nakimbugwe (Co-PI from Nutreal Ltd)
- Ms. Lovin Kobusingye (Co-PI from Kati Farms Ltd)
- Dr. Robinson Odong (Project Officer, Department of Zoology, Entomology and Fisheries Sciences, Makerere University)
- Ms. Juliet Ogubi Nafula (M&E Expert)
- Ms. Nelly Badaru (Gender Specialist)
- Mr. Eugene Swinnerstone Miheso (IT Expert)
- Mr. Maurice Ssebisubi (Kati Farms Ltd)
- Ms. Hasifa Kabejja (Communication Officer)
- Ms. Eve Akongo Ampa (Project Administrator)

Strengthening MEL and Data Quality
During the training workshop, Ms. Juliet Ogubi Nafula guided the team through the NutriFishPlus MEL framework, highlighting its objectives, indicators, results chain, and reporting requirements. Particular emphasis was placed on the role of high-quality data in facilitating project learning, adaptive management, and accountability to donors and beneficiaries.
A key highlight of the training was a hands-on orientation to the household survey tool, led by Mr. Eugene Swinnerstone Miheso. Participants were taken through the structure of the questionnaire, which covers household demographics, nutrition and dietary diversity, fish consumption, livelihoods, value addition, gender dynamics, and food security. Practical sessions enabled the research assistants to practice administering the tool, apply effective probing techniques, and address common field-level challenges.

The training underscored critical ethical considerations in household data collection, including informed consent, confidentiality, and respectful engagement with respondents. Data quality assurance measures such as accurate data recording, use of digital data collection platforms, and daily review protocols were discussed to ensure consistency and reliability.

In his closing remarks, Dr. Efitre emphasized that well-trained research assistants are central to generating credible evidence that reflects the lived realities of beneficiary households. “This training is a vital investment in strengthening our MEL system and ensuring that the evidence we generate informs learning, accountability, and improved project outcomes,” he noted.

The participants expressed gratitude for the interactive and practical nature of the training, noting that the sessions significantly enhanced their confidence and readiness for fieldwork. The training concluded with a question-and-answer session and a brief assessment to confirm participants’ understanding of the MEL framework and household survey tool.

Pre-testing of the household survey tool
Following the training, the project team conducted a pre-test of the household survey tool at Kasenyi Landing Site in Entebbe on 19th January 2026, ahead of the baseline survey. The baseline data collection will be carried out across five landing sites: Katosi (Mukono District), Kikondo (Buikwe District), Bangaladesh and Kayago (Amolatar District), and Dei (Pakwach District), spanning Lakes Victoria, Kyoga, and Albert.
Details on the project at: https://news.mak.ac.ug/2025/10/mak-conas-launches-nutrifishplus-project/
Natural Sciences
Dr Oruru Bosco Appointed Acting Head of the Department of Physics
Published
4 weeks agoon
January 6, 2026
Dr Oruru Bosco has been appointed Acting Head of the Department of Physics at the College of Natural Sciences (CoNAS), Makerere University effective 1st January 2026. Dr Oruru takes on from Dr Okello Denis who has served in the same position since January 2022. He becomes the 10th Head of Department since its establishment. His predecessors include Prof. David Thomson, Prof. John Odeke Ilukor, Prof. Eldad J.K. Banda, Prof. Ezra K.S. Mugambe, Prof. Yusto Kaahwa, Prof. Ezra M. Twesigomwe, Prof. Florence Mutonyi D’ujanga, Prof. Winston Tumps Ireeta, and Dr. Denis Okello.
Dr. Oruru holds a PhD in Physics from the University of the Free State, South Africa, an MSc in Physics from the University of the Free State, a BSc in Astrophysics and Space Science from the University of Cape Town, a BSc in Education (Physics and Mathematics) from Mbarara University of Science and Technology, a Postgraduate Diploma in Public Administration and Management from Uganda Management Institute, and a Diploma in Secondary Education (Physics and Chemistry) from the Institute of Teacher Education, Kyambogo.

He has served the Department in various capacities as a Lecturer, Senior Lecturer, Graduate Supervisors Trainer, and Secretary of the Departmental Research and Teaching Programmes Committee. Dr. Oruru has supervised and examined several PhD, MSc, and BSc students at Makerere and other institutions, including Kyambogo University, Mbarara University of Science and Technology, the South African Institute of Physics, and the University of the Free State. His research interests include accretion onto magnetized compact objects (e.g., X-ray binaries and cataclysmic variables), exotic phenomena such as gamma-ray bursts, supernovae explosions, and active galactic nuclei, African cultural astronomy, and radiation physics.

At the handover ceremony held today, 6th January 2026, Dr. Denis Okello congratulated Dr. Oruru on his appointment and expressed gratitude to the College and University administration for the support accorded to him during his tenure. He also thanked the department staff for their cooperation and commitment. “The support I received enabled me to serve successfully and build upon the achievements of my predecessors. I leave the Department in a much improved state, with well-established infrastructure, including a lecture theatre, classrooms, teaching, research, and ICT laboratories, as well as a departmental library stocked with textbooks relevant to our programmes.”

During his tenure, Dr. Okello oversaw the review of the Bachelor of Science programmes (BSCI and BSCX) to align with the competence-based curriculum. The Department also acquired 55 computers through government funding, in addition to three iPads donated through the College. Renovations were carried out on the interior and exterior of the Department. Staffing improved, with 17 teaching staff currently in place out of the approved 27 positions. Recruitment efforts continue to fill the vacant positions.

Dr Okello appealed to staff to continue working together to further strengthen the department, and pledged support to the new head.
In his remarks, Dr Oruru thanked the staff and university management for entrusting him with the leadership of the Department. ““I am delighted to work alongside a team of dedicated professionals who understand their responsibilities and remain focused on our shared goals. I step into this position not merely as a leader, but as a representative of an exceptional team. With the guidance and support of my predecessors, I am confident that we will continue to uphold and build upon the department’s tradition of excellence.

The Dean, School of Physical Sciences, Dr Michael Owor commended both outgoing and incoming Heads for their commitment to the growth and success of the Department.




Natural Sciences
Jinja Fish Festival: Mak Recognized for its outstanding contribution to the Development of the Fisheries Sector
Published
2 months agoon
December 8, 2025
Makerere University, through the College of Natural Sciences (CoNAS), has been recognized for its outstanding contribution to the development of the fisheries sector in Uganda. The award was presented during the Sustainable Fisheries, Aquaculture, and Environmental Awards Ceremony at the 7th edition of the Jinja Fish Festival, held on 6th–7th December 2025 at Across the Nile Resort, along the banks of the River Nile in Njeru Municipality.
Renowned for showcasing diverse fish cuisines and cultural exhibitions, this year’s festival aimed to promote knowledge exchange, raise awareness, and highlight recent developments around Lake Victoria.

Launched in 2018, the Jinja Fish Festival has become a key platform for the public and stakeholders to learn about sustainable fisheries and the vital role of fish in livelihoods and nutrition. It also provides an avenue for fisheries stakeholders to deliberate on strategies to improve and sustain the sector.
The event includes the Lake Victoria Marathon, Nalubaale boat rowing competitions, a high-level fisheries forum, workshops, and exhibitions. It is organized by the Lake Victoria Fisheries Organization (LVFO) in partnership with the Directorate of Fisheries Resources, Sustainable Fisheries Initiative (SFI), Uganda Fish Processors and Exporters Association (UFPEA), and the International Fish and National Organizing Committee. It is funded by GIZ–SAF Project, the Food and Agriculture Organization (FAO), the International Fund for Agricultural Development (IFAD), and Enabel.

National Fisheries and Aquaculture Forum
A key highlight of the festival was the National Fisheries and Aquaculture Forum, themed “Promoting Sustainable Fisheries and Aquaculture through Ethical and Inclusive Blue Economy Practices.” The forum featured panel discussions on:
- Aligning Uganda’s Blue Economy with global and regional fisheries management frameworks.
- Adoption of climate-smart technologies in fisheries and aquaculture.
- Integration of business strategies and human rights principles into the fisheries and aquaculture value chain.
- Sustainable development of the fisheries and aquaculture sector.

Participants emphasized the importance of strengthening value chain governance, scaling up innovations, and aligning initiatives with youth employment opportunities.
Delivering the keynote address, Mr. Richard Ogutu-Ohwayo, Coordinator at African Lakes Network and Independent Researcher, highlighted the urgent need to address “dirty fish” corruption, improve fisheries resource management through effective policies, and increase human resource capacity within the sector. Dr. Odongokara Konstantine, a Research and Policy Expert, emphasized domesticating international policies into Uganda’s aquaculture practices and building stakeholders’ capacity to access regional and global opportunities.

NutriFishPlus Project Participation in the Festival
The NutriFishPlus Project team, along with students from CoNAS participated in the 7th edition of the Jinja Fish Festival on 6th and 7th December 2025. Led by Ms. Nelly Badaru, Gender Expert on the Project and Ms. Juliet Ogubi Nafula, PhD student in the Department of Zoology, Entomology, and Fisheries Sciences at CoNAS, the team showcased products developed under the project and educated participants on the nutritional benefits of small pelagic fishes.
In its first phase, titled NutriFish, Makerere University researchers, led by Dr. Jackson Efitre, in collaboration with the National Fisheries Resources Research Institute (NaFIRRI – NARO), NUTREAL Limited, and McGill University, Canada, developed five high-value nutritious products, including baby food, sauce, maize meal, snacks, and seasoning, using Mukene (silverfish). The team also published a Mukene recipe book, containing 16 recipes rich in protein, calcium, zinc, and iron.

Through the project, fishers gained access to loans for purchasing boats and fish containers that improve hygiene and product quality. Two enterprise fishing groups, with over 70 boats, were established to ensure consistent supply and standards. Researchers also developed the eCAS system, an application for tracking fish catches. By September 2022, 8,960 catch records had been transmitted through the system, which is now used by over 200 stakeholders, including the Directorate of Fisheries Resources and the Fisheries Protection Unit, helping prevent overfishing.

Other achievements registered under the project included the development of solar tent driers – greenhouse-like structures that reduce the risk of contamination and provide clean and efficient storage and drying, especially during the wet season subsequently reducing post-harvest losses. The project conducted awareness training for 326 people (200 women) to reduce gender inequalities. As a result, domestic violence dropped by 30%, and the number of women in the fish value chain increased. The project also trained eight Graduate students, who have gained valuable skills and knowledge thus boosting critical human resource needs in the fisheries sector.

Other exhibitors at the festival included (NaFIRRI – NARO), FAO, INNOECOFOOD Project at Makerere University, the International University of East Africa, Busitema University, SEETA University (Science, Environment, Engineering, Technology, and Agriculture), Yalelo Uganda, and Preventive Care International, all showcasing emerging technologies and innovations in fisheries.

NutriFishPlus Phase II
Funded by the International Development Research Centre (IDRC) and the Australian Centre for International Agricultural Research (ACIAR) under CultiAF2, the NutriFishPlus Project aims to scale up Phase I activities to further enhance incomes and livelihoods of Uganda’s fishing communities.
Launched on 28th October 2025, the 32-month project will run until March 2028. Its objectives include:
- Scaling up sustainable fishing technologies, including solar tent driers and raised racks, to reach new communities.
- Strengthening market access and supply chain linkages for high-quality fish and fish-based products.
- Empowering women and vulnerable groups through diversified income-generating activities.

Implemented by the Department of Zoology, Entomology, and Fisheries Sciences at Makerere University, in collaboration with Nutreal Ltd and Kati Farms Ltd, the project is led by Dr. Jackson Efitre (Principal Investigator).
Expected outcomes include:
- Improved livelihoods and incomes for marginalized fishing groups, especially women and youth.
- Better health and nutrition through diversified, market-driven fish products.
- Sustainable fish processing and marketing models replicable across Uganda and the East African region.
- Enhanced participation of women and youth in decision-making and benefit-sharing within the Small Pelagic Fishes (SPFs) value chain.
- Strengthened community resilience and ecosystem health through participatory approaches.

Project activities will be conducted at five landing sites around Lakes Victoria, Kyoga, and Albert: Katosi and Kikondo (Mukono and Buikwe districts), Bangaladesh and Kayago (Amolatar District), and Dei (Pakwach District).




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