Global Food Security 33 (2022) 100618
Contents lists available at ScienceDirect 
Global Food Security 
journal homepage: www.elsevier.com/locate/gfs 
Fruit and vegetable biodiversity for nutritionally diverse diets: Challenges, 
opportunities, and knowledge gaps 
Jody Harris a,*,1, Maarten van Zonneveld b,1, Enoch G. Achigan-Dako c, Babar Bajwa d, 
Inge D. Brouwer e, Dhrupad Choudhury f, Ilse de Jager e, Bart de Steenhuijsen Piters e, 
M. Ehsan Dulloo g, Luigi Guarino h, Roeland Kindt i, Sean Mayes j, Stepha McMullin i, 
Marcela Quintero k, Pepijn Schreinemachers a 
a World Vegetable Center, Thailand 
b World Vegetable Center, Taiwan 
c Laboratory of Genetics, Biotechnology and Seed Sciences, University of Abomey-Calavi, Benin 
d CABI, Pakistan 
e Wageningen University and Research, Netherlands 
f International Centre for Integrated Mountain Development (ICIMOD), Nepal 
g Alliance of Bioversity International and CIAT, Mauritius 
h Crop Trust, Germany 
i World Agroforestry, CIFOR-ICRAF, Kenya 
j Crops For the Future, UK 
k Alliance of Bioversity International and CIAT, Colombia   
A R T I C L E  I N F O   A B S T R A C T   
Keywords: Planetary health brings together intrinsically linked issues of human health and natural systems. This paper 
Plant genetic resources reviews evidence of how agrobiodiversity underpins dietary diversity for current human populations in the 
Agrobiodiversity context of fruits and vegetables, and ways to maintain and improve these for future generations. 
Nutrition Both the conservation and sustainable use of fruit and vegetable biodiversity and the consumption of diverse 
Food systems 
Policy diets are sub-optimal, and in many contexts getting worse. Agrobiodiversity and nutrition are linked through 
Evidence review food availability, access, conservation and consumption, with potential win-wins but notable trade-offs for policy 
and action through time, place, agrobiodiversity use, and equity. We pinpoint research gaps and call for inclusive 
deliberation for action.   
1. Background vegetable biodiversity towards our dietary preferences, which have 
co-evolved with the availability of different fruits and vegetables in 
The idea of planetary health brings together intrinsically linked is- different agroecological settings. The use and maintenance of this bio-
sues of human health and the state of the natural systems on which we cultural heritage in contemporary food systems depends on a wide range 
depend (Dangour et al., 2017). Within this framework, it has long been of social, cultural, political, environmental, and economic factors, and 
understood that nutrition and biodiversity are connected (Johns and there is an important interplay between current and future availability 
Eyzaguirre, 2006). Fruits and vegetables are an important part of diets – of fruit and vegetable biodiversity, and current and future dietary di-
providing essential nutrients, phytonutrients, and fibre for health (FAO, versity, within these complex contexts. 
2021) – and of agrobiodiversity – about 1,100 vegetable species are This paper reviews literature produced in the fifteen years since 
recognized worldwide (Meldrum et al., 2018) and there are at least 1, Johns and Eyzaguirre’s initial study on the issue of biodiversity and 
250 documented fruit species in Latin America alone (Bioversity Inter- nutrition more broadly (Johns and Eyzaguirre, 2006). We focus specif-
national, 2021). Over millennia, human activities have shaped fruit and ically on recent evidence of how fruit and vegetable biodiversity 
* Corresponding author. 
E-mail address: jody.harris@worldveg.org (J. Harris).   
1 Lead authors. 
https://doi.org/10.1016/j.gfs.2022.100618 
Received 17 December 2021; Received in revised form 25 January 2022; Accepted 29 January 2022   
Available online 12 April 2022
2211-9124/© 2022 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
J. Harris et al.                                                                                                                                                                                                        G  l o  b a  l  F  o  o d   S  e c  u  r i ty 33 (2022) 100618
underpins dietary diversity for current populations, and evidence on genetic losses in fruit and vegetable species (Khoury et al., 2021). For 
ways to conserve and improve these for future generations. The aim is to some crops, such as tomato, farmers have already replaced most local 
bring together key research findings to frame what we know about links varieties in many regions (Cebolla-Cornejo et al., 2012; Walters et al., 
between fruit and vegetable biodiversity and dietary diversity. A food 2018) and the development of new varieties relies almost entirely on the 
systems lens – encompassing all the elements and activities that relate to planting materials safeguarded in crop genebanks and seed-saver net-
producing, transforming, delivering, and consuming food for humans – works (Bauchet and Causse, 2012). But whereas tomato is relatively well 
is used to better understand the links between healthy people and a represented in existing genebank collections, the genetic resources of 
healthy planet (HLPE, 2017, 2020), and identify challenges and op- most other fruits and vegetables are poorly conserved in genebanks, or 
portunities; knowledge gaps for further research; and support for not at all. For example, 39% of 883 globally-assessed wild fruit and 
decision-makers and practitioners in achieving global development vegetable species requires urgent conservation because they are poorly 
goals. or not conserved in genebanks or in protected areas; another 58% has a 
medium priority for conservation; while only 3% is well conserved 
2. Changes and challenges (Khoury et al., 2019). A quarter of the 1,100 recognized vegetable 
species worldwide have no samples at all conserved in any genebank 
2.1. Threats to fruit and vegetable biodiversity (Meldrum et al., 2018), and most fruit tree species and their wild rela-
tives, particularly those of tropical origins, are challenging to conserve 
While overall biodiversity is defined as the sum of all living organ- in genebanks because their seed does not tolerate the desiccation and 
isms at the genetic, species, and ecosystem levels, agrobiodiversity is low temperatures of orthodox seed storage, and most fruit cultivars have 
more narrowly defined as the range of species, varieties, and ecosystems specific genetic combinations that can be maintained only through 
- together with crop wild relatives, pollinators, and other associated vegetative propagation (Dawson et al., 2013). Without better conser-
organisms - which are used by humans for food and agriculture (FAO, vation options on farms, at landscape level, and in protected areas, and 
2004; Frison et al., 2011). Agrobiodiversity sustains and stabilizes both without genebank back-up, these fruit and vegetable genetic resources 
agro and natural ecosystems of food production and harvesting, and is are at risk of being lost as land use, climate, and agriculture change. 
the source of genetic diversity for developing future foods that can 
tolerate changing environments and keep pace with changing food 2.2. Changing dietary diversity and fruits and vegetables 
needs and preferences. 
Fruit and vegetable biodiversity is part of agrobiodiversity. For the Dietary diversity is defined as the variety of foods consumed in a 
purpose of this review, this is defined as any genetic planting material of certain time period, and thus far has been described for children, 
fruit and vegetable species, and their wild relatives, for current and women, and households, with higher individual dietary diversity asso-
future food and agriculture, in line with the concept of Plant Genetic ciated with better nutrient adequacy (Arimond and Ruel, 2004; Arimond 
Resources for Food and Agriculture (PGRFA). Besides the large number et al., 2010). Therefore, from a food system perspective, diversifying 
of vegetable and fruit species that occur worldwide, crop wild relatives diets both within and across food groups is a key strategy. Consuming a 
are a special group of genetically-related species of fruit and vegetable diversity of fruits and vegetables—with the nutrients, phytonutrients 
species, which can be used in breeding new varieties because they can and dietary fibres contained therein—is important for good health (FAO, 
provide characteristics related to climate resilience and other desirable 2020). Studies have suggested intake ranges of 300–600g per day 
traits (Kilian et al., 2021). Below the species level, local fruit and (200–600g of vegetables and 100–300g of fruits) to meet the twin goals 
vegetable varieties and wild populations of fruit and vegetables are also of human health and sustainable food supply without overly negative 
part of a biocultural heritage with unique tastes and histories (Dwivedi effects on the environment (particularly through water use and chemical 
et al., 2019; van Zonneveld et al., 2018). These provide for distinct food inputs) (Afshin et al., 2019; Loken et al., 2020; Willett et al., 2019). The 
cultures, as well as an important source of genetic variation needed for World Health Organisation (WHO) recommends adults to eat at least 5 
developing new cultivated varieties with traits to ensure current and portions or 400g of fruits and vegetables per day for protective nutri-
future food supplies (Jansen et al., 2020; Kilian et al., 2021; Schouten tional effects (World Health Organisation, 2003), with national 
et al., 2019). food-based dietary guidelines translating these into recommendations to 
Contemporary fruit and vegetable biodiversity continues to decline eat multiple portions of a variety of fruits and vegetables each day 
in farmers’ fields, at landscape level, and generally in ecosystems, in (Herforth et al., 2019). Despite this need for diversity, intraspecific fruit 
parallel with the rapid global decline in overall biodiversity (Díaz et al., and vegetable biodiversity, and diversity of diets within the fruit and 
2019). Ecosystems in 88% of the world’s 846 terrestrial ecoregions are vegetable food group in particular, are little explored in nutrition, which 
poorly conserved, degraded, or disappearing as a result of human ac- tends to look for diversity across food groups only (World Health Or-
tions (Dinerstein et al., 2017). The richness and abundance of wild fruit ganization, 2015). 
and vegetable species, as well as the crop wild relatives of fruit and Globally, the intake of fruit and vegetables remains low for a ma-
vegetable species and pollinators and seed dispersers, decline with the jority of the population (Afshin et al., 2019; Kalmpourtzidou et al., 
degradation and loss of these ecosystems under the pressures of land-use 2020). Low fruit and vegetable consumption is among the top five risk 
change, global climate change, and other threats (Díaz et al., 2019; factors for poor health, with over 2 million deaths and 65 million 
Pilling et al., 2020). Not all crop wild relatives are threatened, and Disability-Adjusted Life Years (DALYs) attributable to low intake of 
several benefit from environmental degradation when they become fruits, and 1.5 million deaths and 34 million DALYs attributable to low 
weeds or invasive (Syfert et al., 2016). But endemic and other intake of vegetables worldwide each year, and particularly in low- and 
narrowly-distributed wild relatives are the most vulnerable to extinction middle-income countries (Afshin et al., 2019). Low consumption of 
caused by land use change and degradation following the criteria of the fruits and vegetables is a global problem: only 7% of countries in Africa, 
International Union for Conservation of Nature’s (IUCN) Red List of 7% in the Americas, and 11% in Europe reach 240 g/day of vegetables 
Threatened Species (Cadima et al., 2014; Khoury et al., 2020; Schelde- per person on average (Kalmpourtzidou et al., 2020). Looking at 
man et al., 2006). within-country variation, only 20% of individuals in low- and 
Four out of five studies on crop genetic erosion has found evidence of middle-income countries reach the recommendation of 5 servings of 
crop diversity loss, the magnitude varying by species, geographic scale, fruits and vegetables per day (Frank et al., 2019). 
and region, as well as analytical approach (Khoury et al., 2021). So far, The nutrition transition, whereby global diets are moving away from 
most genetic erosion studies have been done on cereal crops and their local traditional patterns through changing demographic and food sys-
wild relatives; few studies are available on the rate of varietal and tem drivers, may make diverse fruits and vegetables either more or less 
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locally available, depending on context (Global Panel on Agriculture and at least in part on foods gathered in their surrounding environment 
Food Systems for Nutrition, 2016; Popkin et al., 2020). While global (World Health Organization, 2015). In some contexts analysed, up to 
fruit and vegetable supply has started to increase proportional to staple 75% of consumed vegetables and fruits are gathered rather than culti-
crops in recent decades, global focus remains predominantly on cereal vated or bought (Powell et al., 2015). 
production (Gould, 2017), and documented supplies remain insufficient Despite the wide occurrence of food gathering, it has been suggested 
for most countries to meet the WHO recommendation: in 1965, suffi- that reliance on collecting wild food plants is often a coping mechanism 
cient fruits and vegetables (≥400 g/day) were available for 17% of the in the face of scarcity, rather than an active choice, which needs to be 
global population, increasing to 55% in 2015 (Mason-D’Croz et al., better understood (Jones, 2017). In addition, overharvesting is a threat 
2019). Supply varies widely: in Africa, only 13% of countries have an to the genetic resources of some wild-harvested species, and high de-
adequate vegetable supply, while in Asia 61% do (Kalmpourtzidou et al., mand for wild food plants could reinforce this threat, or even deteriorate 
2020). All of these data are limited, likely missing the proportion of whole ecosystems. For example, in the case of buruti, an ecologically 
neglected and underutilized and wild-harvested food species, missing important palm species from the Amazon that is valued for its vitamin-A 
food loss and waste, and missing within-country variation. In Africa rich fruits, people destructively harvest female plants, degrading wild 
particularly, the consumption of wild vegetables is reported in many buruti populations with potentially negative impacts on species richness 
countries (Achigan-Dako et al., 2011; Maundu et al., 2009), but data are and carbon stocks of Amazonian peatlands (Bhomia et al., 2019; Endress 
unavailable on consumption frequency or amount across the continent, et al., 2013). Similar cases of overharvesting have been observed around 
and the level of wild harvest is rarely documented. the world, and there has been call to domesticate threatened food plants 
Fruits and vegetables purchased through markets are unaffordable so that people can grow them in their yards or farm fields, to reduce 
for many, with three billion people, and up to 90% of the population in harvesting pressure on wild populations while making these nutritious 
some countries, unable to afford diverse healthy diets in 2017 (Herforth foods available and accessible to a wider public (Akinnifesi et al., 2006; 
et al., 2020). Fruits and vegetables appear more affordable than staple Sundriyal and Sundriyal, 2003). In parallel, research and development 
foods when comparing prices per micronutrient, where they are likely to initiatives are supporting decision-makers and communities in the 
be a relatively low-cost source of varied vitamins, minerals, and phy- implementation and monitoring of sustainable fruit and vegetable har-
tonutrients – but this is not how most families choose their food, with vesting practices at landscape level (Van Loon et al., 2021). 
particularly poorer households focusing on acquiring calories. Even if While agrobiodiversity may have only moderate associations with 
fruits and vegetables are available, accessible and affordable, most dietary diversity at the household level, availability of a diversity of 
people still do not consume sufficient quantities (Hall et al., 2009), fruits and vegetables is a prerequisite for diverse diets at the food system 
particularly if they are not considered an acceptable or desirable food level. As a sum of local changes, in food systems globally food plant 
choice, for instance due to food safety concerns, taste, convenience, or diversity is associated with a food system’s capacity to provide sufficient 
cultural appropriateness, or if people have low knowledge or awareness nutrients through different environmental, social, and economic shocks 
about the importance of fruit and vegetables for health (Aggarwal et al., (including population growth and climate change), but over the past five 
2016; Ha et al., 2020; Hammelman and Hayes-Conroy, 2014),. decades this capacity has declined in all regions except Asia (Nicholson 
et al., 2021; Pingali, 2012). Studies have found that existing agro-
2.3. Bringing fruit and vegetable biodiversity and dietary diversity biodiversity is underutilized in national food systems, with species 
together consumption globally moderate, and lowest in low-income countries 
(Jones et al., 2021) and many food species limited to consumption in 
Food availability, accessibility, affordability, and desirability are small geographic regions (Lachat et al., 2018) - though caveats on the 
therefore the food system links between agrobiodiversity and dietary limitations of data on wild and local foods, discussed above, still apply. 
diversity (Toledo and Burlingame, 2006; Turner et al., 2018) with more The current food system in many ways contributes to agro-
species-diverse food associated with better micronutrient-quality diets biodiversity loss (Hunter et al., 2016), and fruit species and to a lesser 
(Lachat et al., 2018). In specific agricultural diversity studies, and degree vegetables are among the major crop types dependent on polli-
mainly in rural areas, agrobiodiversity has been associated with dietary nation services that are in decline (Gallai et al., 2009). Changing diets 
diversity at the local level: greater crop diversity managed by farming globally as a result of socio-economic, demographic, agricultural, and 
households is usually associated with greater dietary diversity at the food system pressures is a process that while it can bring opportunities of 
farm household level, though depending significantly on local agro- scale and trade, can also be seen as a narrowing of the food base on 
ecological context(Jones, 2017). Associations between crop diversity which humans depend (Frison et al., 2006; Powell et al., 2013; World 
and fruit and vegetable intake have been seen specifically in some Health Organization, 2015). Large structural changes such as global-
studies (Herforth, 2010; Jones et al., 2014), though many do not ization of supply chains and societies, and changing demographics and 
examine this connection explicitly, and fruits and vegetables are often urbanisation, have shaped food regimes over the past half-century to 
combined into a single food group or assessed together with other plant prioritise foods that are less perishable and more globally tradable (Lang 
foods such as pulses. Magnitudes of association tend to be very small and Heasman, 2015; Magnan, 2012), side-lining perishable fruits and 
when only factoring in cultivated foods (Jones, 2017), prompting some vegetables requiring local production, processing, or complex food 
to suggest that increasing production diversity is not an efficient tool to chains to get them safely from farm to fork. Large-scale, high-input 
improve nutrition at the household level (Sibhatu and Qaim, 2018). monoculture production systems for pineapple, banana, and avocado 
Associations may be stronger when including wild food plants in among other commercial fruit crops, have led to excessive environ-
agrobiodiversity metrics in contexts where these are eaten: In Kenya, for mental degradation and biodiversity loss (Magrach and Sanz, 2020; 
example, when food plants collected from a wider landscape are taken Ploetz, 2021; Shaver et al., 2015). In the case of banana, genetic 
into account, a one unit increase in household access to agrobiodiversity impoverishment has led to a highly vulnerable supply system that 
(species diversity score) is associated with a 13% increase in probability largely relies on a single clone that is susceptible to Panama disease, 
of micronutrient adequacy in the diet (Oduor et al., 2019). In a similar threatening global banana supply (Kema et al., 2021; Ploetz, 2021). 
way, a case study in Tanzania found a significant association between In addition to the micronutrients in fruit and vegetable species that 
diversity of vegetables produced or collected from the wider landscape, we know about, there are also a range of phytonutrients and bioactive 
and micronutrient adequacy and dietary diversity (Keding et al., 2012). compounds across fruit and vegetable species, whose nutritional func-
In some settings and seasons, wild-harvested fruits and vegetables tions are still poorly understood (Lutaladio et al., 2010). Despite issues 
contribute significantly to nutrient intake (Powell et al., 2013) and in- of bioavailability of nutrients and the various effects of processing and 
comes (McMullin et al., 2021), with estimates of a billion people relying cooking on available nutrients, dietary diversity within fruits and 
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vegetables is likely to be important, and conserving fruit and vegetable genebank network are vulnerable because they have large backlogs of 
biodiversity for current and future availability may protect health old or original fruit and vegetable germplasm samples that require 
through food in ways we don’t yet know. propagation and re-storage (Fu, 2017). 
Complex access and benefit sharing policies and regulations, in 
3. Options and actions particular domestic policies and regulations that implement the Nagoya 
Protocol of the Convention on Biological Diversity (CBD), increasingly 
3.1. Maintaining fruit and vegetable biodiversity for current and future govern international efforts to conserve and use the diversity of local 
generations varieties and wild populations (Brink and van Hintum, 2020). These 
policies recognize the rights of countries and local communities over 
Several important trends are increasing the conservation and use of genetic resources within their territories, yet all countries and commu-
fruit and vegetable biodiversity in food systems at global and local nities depend on each other for genetic resources of fruit and vegetable 
levels. In addition to the fact that the proportion of fruit and vegetables species for food and nutrition, including neglected and underutilized 
in global food production is increasing (Gould, 2017; Khoury et al., species (Khoury et al., 2016; van Zonneveld et al., 2021). This interde-
2014; Martin et al., 2019), advanced technologies are now accessible to pendence is expected to increase under global climate change (Burke 
public and private breeders and researchers globally to mainstream the et al., 2009), and regulations will need to evolve alongside. 
use of genetic diversity for developing new varieties of fruits and veg- Currently, germplasm exchange of most fruit and vegetable species is 
etables (Jamnadass et al., 2020; Schouten et al., 2019). Cities are limited, including because they are underrepresented in the Multilateral 
becoming important hubs of crop diversity because immigrants bring System of the International Treaty on Plant Genetic Resources for Food 
planting material from their home areas (Rimlinger et al., 2021; Taylor and Agriculture (Plant Treaty) that is established to enhance exchange of 
and Lovell, 2014), while at the same time the coverage of protected plant genetic resources for food and agriculture between countries. 
natural and biocultural environments has tripled in the last 40 years Furthermore, debates about the relationship and interplay between 
(Pringle, 2017), and at least 35% of the terrestrial protected areas are breeders’ rights and farmers’ rights are ongoing (Dias, 2011; Gupta and 
owned and/or managed by local and Indigenous communities, who play Negi, 2019; Salazar et al., 2007), affecting the development of new plant 
an important role in maintaining agrobiodiversity worldwide (Díaz material and the use of seed by growers. The 2011–2021 Global Crop 
et al., 2019). Although these trends could possibly bend the curve of Wild Relatives project led by the Crop Trust showed how global part-
decline in fruit and vegetable biodiversity, they may not completely nerships for collecting, conservation, and germplasm availability are 
halt, let alone reverse, the loss. For example, the expansion of protected possible for crop wild relatives of fruit and vegetable species that fall 
areas provides some opportunities for conservation, but ecosystems in under the framework of the Plant Treaty, such as banana, apple, 
these areas may be degraded already, and fruit and vegetable biodi- eggplant, and carrot (Müller et al., 2021). Similar workable agreements 
versity within the landscape may decline (Pringle, 2017). enable rescue of germplasm of other fruit and vegetable species and give 
Crop wild relatives stand further away from people’s diets and are farmers, breeders, and researchers access to planting material for food 
often relevant for breeding only. Their conservation is taken up by many and nutrition; these agreements provide a framework for access and 
governments in protected area management (Dulloo and Maxted, 2019), benefit sharing following all applicable current laws and regulations at 
but crop wild relatives of fruit and vegetable species are still of low national and international level (Brink and van Hintum, 2020). 
priority compared to those of cereals and pulses; it is important to raise 
awareness about the need to conserve the former as well. 3.2. Enabling access to a wide range of fruits and vegetables 
Local fruit and vegetable species and varieties are still maintained 
and shared by farmers and communities in different production systems Promoting neglected and underutilized food plants, such as many 
(Dulloo et al., 2017). These plants provide nutritional and food security, traditional fruit and vegetable species, is a key policy option that ad-
income-generating opportunities, and ecosystem services, and dresses both healthy diets and agrobiodiversity (Pedersen et al., 2020). 
contribute to cultural identity (Sthapit et al., 2016), and also represent a The pool of underutilized plant biodiversity includes fruit and vegetable 
valuable conservation of biodiversity for food. To preserve these, gov- species with exceptionally high nutritional value, such as 
ernments and societal actors can recognize the custodian farmers and micronutrient-rich African leafy vegetables adapted to rain-fed condi-
communities who maintain these unique and traditional production tions (Maundu et al., 2009) and vitamin-rich Amazonian fruit trees that 
systems, such as governments in some countries are doing already for withstand flooding and waterlogging (Van Loon et al., 2021; van Zon-
important agricultural heritage systems (Koohafkan and Altieri, 2011). neveld et al., 2020), giving impetus to connections between biodiversity 
Governments and societal actors can further support these farmers and and nutrition. The latest generation of food-based dietary guidelines 
communities to establish or maintain seed networks, and encourage starts to move in the direction of more diverse diets – but these efforts 
equitable business linkages to markets for more resilient livelihoods can better consider cultural acceptability, and may require promotional 
based in maintaining biodiversity, through incentives and regulation efforts to increase the willingness of consumers to shift their diets to new 
(Dulloo et al., 2017). or forgotten foods and diversify within the fruit and vegetables food 
Complementary to local conservation on farms and in protected groups (Davis et al., 2021). Alongside improving availability and access, 
areas and local seed-sharing, large national and international fruit and the challenge is to enhance consumer choice of, and preference for, these 
vegetable germplasm collections have been established in genebanks in foods. There is clear evidence that focusing on education at all levels is 
North America, South America, Asia, and Europe (Byrne et al., 2018; to some extent effective for modifying behavioural changes in general, 
Cunha Alves and Azevedo, 2018; Engle and Faustino, 2007; Jacob et al., whether dietary or environmental (Alderman and Headey, 2017); and 
2015; Loskutov, 2020; Van Den Houwe et al., 2020). They safeguard nutrition literacy, social norms for healthy eating, and self-efficacy are 
fruit and vegetable biodiversity to improve availability of planting ma- key components of health-related behaviour change (Eker et al., 2019). 
terial for food and nutrition of current and future human generations, We know less for fruits and vegetables in particular, though the 
and are a source for breeding and germplasm exchange to develop new general evidence on dietary behaviour change can be tested, and social 
foods for new tastes or new agricultural conditions. Sub-Saharan Africa marketing has been used to promote a return to diverse traditional 
presents a gap in the genebank network; only one out of three countries vegetables in several contexts (Powell et al., 2015). Nutrition literacy 
in this region has a national crop genebank (World Bank, 2017). In- programs generally target women, who are in many cases custodians of 
vestment in genebank infrastructure in this region would help to household nutrition and of local agrobiodiversity, but there will also be 
maintain and document sub-Saharan African fruit and vegetable genetic a need for community-targeted messages to change social norms around 
resources. At the same time, several collections from the existing both diets and conservation (Van den Bold et al., 2013). There are 
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reasons that changing food demand might help to reverse the trends needed for human populations to meet dietary recommendations, but 
reducing agrobiodiversity and homogenizing food systems: There is there are different ways to achieve this, with implications for the use of 
greater global awareness about the benefits of diverse diets with suffi- crop biodiversity at different geographic scales. For sustainable use of 
cient fruits and vegetables, though this knowledge has not reached all fruit and vegetable biodiversity a wide range of approaches depending 
populations equally. Encouragingly, some neglected and underutilized on contexts, including sustainable intensification and diversification 
fruit and vegetable species have regained relevance in urban diets approaches to improve yields while adapting climate change, have been 
through public and private initiatives in gastronomy and niche markets suggested to maintain fruit and vegetable production and livelihoods 
for local, healthy, or ethnic food (Borelli et al., 2020), and appealing to while minimizing environmental degradation (Godfray and Garnett, 
aspiration can be a powerful promoter of foods. But decades of work on 2014; Schreinemachers et al., 2018; van Zonneveld et al., 2020). 
the nutrition transition makes it clear that these positive food system Participatory approaches that consider multiple criteria including 
and dietary changes are the exception rather than the norm (Popkin agronomic, environmental, social, economic, and political factors have 
et al., 2020). prooved useful to guide decision-making in balancing sometimes 
Deciding what to eat is embedded in the broader food environment, competing interests in making these production decisions (van Zonne-
and within people’s daily realities (Blake et al., 2021). Promoting home veld et al., 2020). 
and community fruit and vegetable production, or facilitation of Producing sufficient fruits and vegetables can have relatively high 
foraging, is a small-scale option where the agrobiodiversity and broader requirements for land, water, and chemical inputs, depending on 
context allows (Baliki et al., 2019; Powell et al., 2015; Schreinemachers context and production method (Aleksandrowicz et al., 2016), and with 
et al., 2016), but approaches are needed that can clearly provide for the one third of global greenhouse gas emissions produced by the food 
nutritional needs of large (often urban) populations. Addressing system there is a need to better understand the role of fruits and vege-
affordability of fruits and vegetables is key to creating an environment tables in land and input use (Crippa et al., 2021). While we can in theory 
where all can access a healthy diet, given that most people buy at least a produce sufficient fruits and vegetables for healthy diets within plane-
proportion of their food, and affordability can come from a combination tary boundaries (Willett et al., 2019), achieving national food-based 
of lower retail prices of diverse fruits and vegetables (through produc- dietary guidelines has been found to be incompatible with climate and 
tivity improvements, reduced postharvest losses, or increased market environmental targets in a majority of 85 countries studied (Springmann 
efficiency for stable prices) and higher incomes (from inclusive eco- et al., 2020). One key solution is a better use of fruit and vegetable 
nomic growth, living wages, and social safety nets) (Hirvonen et al., biodiversity in breeding globally to make varieties available to pro-
2019). Cheap food is not necessarily good for healthy diets, fair liveli- ducers that are nutrient-rich, eco-efficient in production, and adapted to 
hoods or agrobiodiversity however, so a focus on affordability through changing climate conditions. Producing according to context is another 
equitable economies is important where foods are purchased (Benton key solution. Local production, home garden production, and food 
et al., 2021), and decisions will have to be made regarding priorities and gathering may be important sources of biodiverse plant foods in many 
trade-offs (below). contexts, to buffer food production homogenization and increase the use 
Big-picture policy and political drivers – including research and of fruit and vegetable diversity at national and local levels (Bharucha 
development investment focus, trade regimes, and a focus on calorie and Pretty, 2010; Galluzzi et al., 2010; Schreinemachers et al., 2015). 
security rather than nutrition – have oriented food systems away from Urban agriculture also brings opportunities for agrobiodiversity in 
pathways prioritising diverse fruits and vegetable species in diets, and urban spaces (World Health Organization, 2015), with benefits 
away from agronomic and food system paradigms that might promote a including reducing food losses along the value chain by producing food 
return to more biodiverse production systems and diverse diets (Leach closer to where it is consumed; creating jobs for urban populations; 
et al., 2020; Patnaik and Oenema, 2015; Rosset and Altieri, 2017; improving quality of life through greening of urban environments; and 
Vivero-Pol et al., 2018). The policy environment in terms of evidence, closing nutrient cycles by re-using urban waste (Lutaladio et al., 2010). 
politics, and capacity – and the framing of these – is important for sus- Agroecology research and practice in particular tries to develop, test, 
tainable change in biodiversity and diets (Hunter et al., 2016), though in and promote practices that promote sustainable production and equi-
global analyses the political commitment is low for conserving and using table diets through inclusive and participatory approaches (Méndez 
fruit and vegetable biodiversity (Jones et al., 2021) and for improving et al., 2015; Rosset and Altieri, 2017), and considers the potential of 
nutrition (te Lintelo and Lakshman, 2015). This is despite efforts such as local fruit and vegetable species for sustainable production in specific 
the Voluntary Guidelines for Mainstreaming Biodiversity into Policies, agroecological zones and adaptation to changing climates (Nabhan 
Programmes and National and Regional Plans of Action on Nutrition, et al., 2020; van Zonneveld et al., 2020; Waha et al., 2018). 
endorsed by the Commission on Genetic Resources for Food and Agri- Current discourses encourage integrated approaches between 
culture (CGRFA) in 2015, which recognize many of these issues but do different conservation practices (Dulloo et al., 2017) and between 
not hold powers of accountability for their action. environmental sustainability and human diets (Willett et al., 2019). 
Notwithstanding, there are trade-offs among conservation of biodiver-
3.3. Synergies and trade-offs sity as an end in itself vs. leveraging fruit and vegetable agrobiodiversity 
to provide food options and improve diets, given scarce resources. The 
There are potential synergies between a food systems approach to approaches mentioned above prioritise agrobiodiversity conservation 
nutrition and an integrated conservation of fruit and vegetable biodi- for human food, which while important for us humans, is not necessarily 
versity on farms, at landscape level, and in protected areas (Powell et al., all that is needed to preserve levels of biodiversity that the planet re-
2013). Linking agrobiodiversity and nutrition can in theory address quires to function well. Within the context of a diet for a healthy planet, 
some of the negative consequences of current food systems (Johns and there are trade-offs and synergies between leveraging agrobiodiversity 
Eyzaguirre, 2006): Promoting diverse food consumption can promote to improve diets with more volume of vegetables and fruits, and making 
conservation of agrobiodiversity; and promoting agrobiodiversity is a food production more eco-efficient with an investment in a broader 
practical approach for dietary diversity, food security, and rural devel- conservation of biodiversity and environmental sustainability. There are 
opment, so there are win-wins to be had in theory (Toledo and Burlin- also trade-offs among our ability to preserve the biodiversity of different 
game, 2006). There are also potential trade-offs between preserving species, and our requirements in diets: Typically, small and orthodox 
agrobiodiversity and enabling diverse diets for everyone everywhere, seed of many vegetables and berries can be saved in genebanks and seed 
alongside promoting fair food system livelihoods, which need to be saver networks for exchange, breeding and future use (Hong et al., 
resolved through evidence, policy engagement, and inclusive dialogue. 1996). In contrast, many fruit species can only be maintained in field 
Clearly, more availability of a variety of fruits and vegetables is collections, or by conserving living tissue in tubes (in vitro), or through 
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cryopreservation in liquid nitrogen because they usually have large and enough understanding of the nuances of different agroecological or 
recalcitrant seed that cannot be stored for long time periods (Panis et al., socio-cultural contexts or production approaches. There are also 
2020). Appropriate planting material is one of the major bottlenecks for research efforts highlighting national-level agrobiodiversity and its use 
farmers to grow a wide range of fruit and vegetable species (McMullin in food systems, and specifically on metrics for measuring this, though 
et al., 2021), so genebanks of vegetable and fruit species can actively little looking at the impacts of national policies on the crossovers 
engage with different users groups to provide inter- and intraspecific identified above. Trading and international interdependence are rele-
crop diversity for developing and growing planting material with high vant topics at regional and global levels, with some literature on broad 
nutrient values for a win-win scenario (Smale and Jamora, 2020; Stoi- food system changes and their impacts on agrobiodiversity and dietary 
lova et al., 2019). The fruit and vegetable biodiversity conserved in diversity, but there is little literature looking at fruits and vegetables at 
genebanks and by seed saver networks complements plant genetic re- this level. Below we highlight key research and knowledge gaps from 
sources maintained in farmer fields, at landscape level, and in protected this review. 
areas (Dulloo et al., 2017). Connecting these genebanks and networks to A key knowledge gap is that we don’t have good data on the diversity 
these efforts on the ground optimizes conservation of fruit and vegetable of fruit and vegetable species in farmer’s fields or landscapes, as well as 
biodiversity while providing local users opportunities to access new and in genebanks, especially within species. There is often high nutrient 
diverse planting material. The latter being particular relevant under variability among fruit and vegetable varieties, a clear link between 
climate change. The way and extent these different practices are agrobiodiversity and nutrition, but data are limited on this, and 
implemented and connected may differ depending contexts, with particularly for neglected and underutilized food plants (Johns and 
different implications for different societal groups in the food system. Eyzaguirre, 2006). These include many traditional fruits and vegetables 
In practice, decision-makers and societal actors may need to make a relevant for local production and harvesting but often left out of data 
choice between the contemporary use of certain species and varieties in collection, policy considerations and agricultural extension (Hunter 
production systems to feed today’s population, and conserving agro- et al., 2019; Raihana et al., 2015). Many locally cultivated and 
biodiversity at a broader scale (including todays’ neglected and wild-harvested food plants are missed in available dietary and produc-
underutilized species and varieties) to safeguard a biocultural heritage tion data (Jones et al., 2021), limiting our knowledge about real diets 
and to keep future food options open. Decisions need to be made about and food environments; at the same time, fruit and vegetable species – 
the levels and type of fruit and vegetable biodiversity to be conserved at which have very different agronomic, trade, and nutritional profiles – 
national and local levels with an eye on the future, depending on the are often lumped together into a single ‘fruits and vegetables’ group, 
local and global importance and potential of certain species and vari- further hampering nuanced research efforts (Harris et al., 2021). In-
eties for human diets under climate change, and in the context of global vestment in screening the diversity of neglected and underutilized fruit 
efforts of conservation and germplasm exchange, such as the Global and vegetable species can reveal varieties with high nutrient content 
System of PGRFA (Engels and Ebert, 2021). Also with an eye on future (Yang and Keding, 2009) providing evidence for nutrition and market-
generations, agrobiodiversity promotion through school gardens, ing messages to promote diet and crop diversification with more fruit 
sometimes in combination with locally-procured school meals, com- and vegetable species among households and producers, and a basis for 
bines early nutrition education and diverse production as a potential breeding of nutrient-rich varieties. 
win-win engaging with future eaters and conservers (Hunter and There is also a key knowledge gap in the diversity of fruit and 
Monville-Oro, 2020; Shrestha et al., 2020). Further down the food sys- vegetable species in people’s diets. There is little research available on 
tem, reduction of food prices through market efficiency and food pro- what is eaten where, in particular the use of wild-harvested food plants 
duction homogenization can lead to agrobiodiversity losses, so a (World Health Organization, 2015), though in some places and in 
trade-off in some contexts is between affordability of food and the different languages there may be ethnobotanical and anthropological 
availability of agrobiodiversity, which requires further understanding literature that can be drawn on more than it currently is – and that 
and debate on agrobiodiversity use in value chains and the amount of rescues and recognizes traditional knowledge from local voices that still 
investment to counteract agrobiodiversity losses through rescue and can be heard in some places on how to grow these species, and how to 
conservation. Considering trade-offs between current and future pop- process them for food consumption. There is also a gap in terms of why 
ulations is part of these deliberations, including with a focus on inter- different fruits and vegetables are preferred in different places, and in 
generational equity (Nisbett et al., 2021). the role of nutritional information, aspiration, and gastronomy in 
Regarding equity for today’s human populations, some of the most inspiring people to eat more diverse plant foods. The lack of data on 
nutrition-insecure people live in some of the most biodiverse areas, so intra-species food composition, and the context-specificity of diets, 
issues of nutrition equity and conservation should be considered in na- means that more research is needed to provide more general policy 
tional and international policies to link fruit and vegetable biodiversity advice (Hunter et al., 2016). 
to dietary diversity (Herrero et al., 2017; Johns and Eyzaguirre, 2006). Much of the research on agrobiodiversity in food systems and for 
For example, while the Amazon is a hotspot of fruit and vegetable diets has focused on rural parts of middle-income countries, where 
biodiversity, these territories are also characterized by complex socio- smallholder farming is common, but other regions are important to 
economic problems that affect local agrobiodiversity as well as health understand. Rural and urban communities in some middle- and high- 
and welfare of indigenous communities, such as the nutrition transition income countries especially in Asia and the Mediterranean region 
and poor health and sanitation infrastructure (Borges et al., 2015; have retained aspects of traditional biodiverse production systems in 
Coimbra et al., 2013). Costs and benefits of conservation and fruit and combination with the maintenance of traditional food consumption 
vegetable production do not accrue equally or fairly in different di- customs, and seem to have better health outcomes compared to those 
mensions of the food system – but different facets of equity have been regions which have not – at least so far (Johns and Eyzaguirre, 2006). To 
described (Nisbett et al., 2021), and can be explicitly factored in to in- which extent biodiverse wild-harvested fruits and vegetables are 
clusive decision-making processes to debate trade-offs and decide pri- accessible for urban dwellers and can be used for food-plant harvesting, 
orities at different levels. also remains to be investigated (Rimlinger et al., 2021; Shackleton et al., 
2017). Low-income countries are particularly under-represented in food 
4. Knowledge gaps system studies of fruits and vegetables for healthy diets (Harris et al., 
2021). 
Overall, there is considerable literature focused on agrobiodiversity We also need better evidence as to how greater agrobiodiversity in 
for local diets, including the role of fruit and vegetable biodiversity in general would bring benefits in terms of healthier diets and more sus-
populated rural areas; but far less on biodiversity in urban areas, and not tainable food production. While raising crop diversity at the level of 
6
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individual farms may indeed not be the most effective strategy to in- Acknowledgements 
crease dietary diversity in those households (Jones, 2017), at the food 
system level producing diversity is of course a necessary precondition The authors wish to thank the United National Food Systems Summit 
for being able to consume a diversity of foods, and we need to better Fruit and Vegetables Working Group for original inputs to the UNFSS 
understand those pathways. There is little research on effective strate- working papers (on vegetable biodiversity and vegetables in dietary 
gies to promote more diversity in modern or informal supply chains diversity) which underpin this work and can be found at https://sc-fss2 
(from producers and traders to processors and vendors), and there is a 021.org/materials/fss-briefs-by-partners-of-scientific-group/. 
need to better understand this missing middle and trade-offs among We thank Gayle Volk for her contribution to these original working 
scaling production for broader availability in diets, and conserving fruit papers. 
and vegetable agrobiodiversity and the broader environment. More The first, second and last authors acknowledge funding support from 
research is needed on the potential of agroecological practices to make long-term strategic donors to the World Vegetable Center: Taiwan, UK 
fruit and vegetable production more compatible with national climate aid from the UK government, United States Agency for International 
and environmental targets. Development (USAID), Australian Centre for International Agricultural 
Given the range of factors important in understanding the links be- Research (ACIAR), Germany, Thailand, Philippines, Korea, and Japan. 
tween agrobiodiversity and dietary diversity – from plant genetic re- The second author received financial support from the German 
sources and ethnobotany, to nutritional screening and breeding, to Federal Ministry for Economic Cooperation and Development (BMZ) 
socio-cultural behaviours and political economy – interdisciplinary commissioned by the Deutsche Gesellschaft für Internationale Zusam-
research is clearly needed (Hunter et al., 2016; Méndez et al., 2015). menarbeit (GIZ) through the Fund International Agricultural Research 
Bringing this research together into a body of evidence can be difficult, (FIA), grant number: 81275070. 
as even standard epidemiological studies of agrobiodiversity and dietary 
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