# Plant proteins for human and environmental health: knowledge, barriers, and levers for their development, a case study in France. A review

Karine Gallardo Guerrero, Anne Saint-Eve, Rémy Cailliatte, Patrick Carré, Guy Della Valle, Jérôme Enjalbert, Mylène Durand-Tardif, Ingrid Lorinet, Claire C. Gaudichon, Jean-Baptiste Guyon, et al.

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## ► To cite this version:

Karine Gallardo Guerrero, Anne Saint-Eve, Rémy Cailliatte, Patrick Carré, Guy Della Valle, et al. Plant proteins for human and environmental health: knowledge, barriers, and levers for their development, a case study in France. A review. Agronomy for Sustainable Development, 2025, 45 (4), pp.42. 10.1007/s13593-025-01034-1. hal-05197858

HAL Id: hal-05197858

https://hal.science/hal-05197858v1

Submitted on 4 Aug 2025

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# Plant proteins for human and environmental health: knowledge, barriers, and levers for their development, a case study in France.
A review

Karine Gallardo-Guerrero<fnref n="1" /> · Anne Saint-Eve<fnref n="2" /> · Rémy Caillatte<fnref n="3" /> · Patrick Carré<fnref n="4" /> · Guy Della Valle<fnref n="5" /> ·

Mylène Durand-Tardif<fnref n="6" /> · Jérôme Enjalbert<fnref n="7" /> · Claire Gaudichon<fnref n="8" /> · Jean-Baptiste Guyon<fnref n="9" />,<fnref n="15" /> · Stéphane Jezequel<fnref n="10" /> ·

Ingrid Lorinet<fnref n="9" /> · Marie-Benoit Magrini<fnref n="11" /> · Catherine Ravel<fnref n="12" /> · Catherine M. G. C. Renard<fnref n="13" />,<fnref n="14" />

Accepted: 16 June 2025

© The Author(s) 2025

## Abstract

Agriculture faces multiple challenges, including reducing negative environmental impacts and producing quality food for a growing world population. One of the significant issues is the need for an increased proportion of plant-based foods in diets in the developed world and its combination with the ecosystemic services provided, such as those provided by legumes, allowing a reduction in the use of nitrogen fertilizers. This requires systemic innovation in order to increase the sustainable production and consumption of diverse plant protein sources in an interaction between knowledge, tools, regulation, and acceptability. The case of France is chosen to reflect on these innovation issues. There have been a number of recent reviews either on production or on food processing, but none has addressed their systemic interrelations, critical to select the levers such as genetics, agronomy, and processing to improve protein quality and use while maintaining crop performance in an environmentally friendly manner. Here, we review the issues of proteins for healthy and sustainable diets in synergy with agronomical benefits and recommend research priorities to address this topic from farm to fork. This review includes (1) analysis of the economic context for plant-based protein production for food in the context of the agroecological transition towards sustainability and on the consequences in plant breeding schemes; (2) assessment of overall protein production from a diversity perspective, including genetics and diversity of the production systems and territories; and (3) plant proteins in healthy and attractive foods: protein content and quality, typology of processing options for the production of protein-rich food products in relation to consumer needs and demands. Next, we identify major knowledge gaps and emphasize the need for transdisciplinary research from plant breeding to the food market, involving the various stakeholders in the food value chain, to address such complex issues for more sustainable agri-food systems.

Karine Gallardo and Anne Saint-Eve share the first authorship.

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Catherine M. G. C. Renard
catherine.renard@inrae.fr

Jean-Baptiste Guyon
jean-baptiste@dlabindustry.com

<footnote n="1">¹ Agroécologie, INRAE, Institut Agro Dijon, Université de Bourgogne, Université de Bourgogne Franche-Comté, 21000 Dijon, France</footnote>

<footnote n="2">² SAYFOOD, INRAE, AgroParisTech, Université Paris-Saclay, 91120 Palaiseau, France</footnote>

<footnote n="3">³ Carnot Plant2Pro, INRAE, 84000 Avignon, France</footnote>

<footnote n="4">⁴ Terres Inovia, 33600 Pessac, France</footnote>

<footnote n="5">⁵ BIA, INRAE, 44000 Nantes, France</footnote>

<footnote n="6">⁶ PlantAlliance, 5 rue Watt, 75013 Paris, France</footnote>

<footnote n="7">⁷ GQE-Le Moulon, Université Paris-Saclay, INRAE, AgroParisTech, CNRS, 91190 Gif-sur-Yvette, France</footnote>

<footnote n="8">⁸ PNCA, INRAE, AgroParisTech, Université Paris-Saclay, 91120 Palaiseau, France</footnote>

<footnote n="9">⁹ Greentech, biopôle Clermont-Limagne, 63360 St Beauzire, France</footnote>

<footnote n="10">¹⁰ ARVALIS - Institut du Végétal, 75016 Paris, France</footnote>

<footnote n="11">¹¹ AGIR, INRAE, INP, Université de Toulouse, 31300 Castanet-Tolosan, France</footnote>

<footnote n="12">¹² Université Clermont-Auvergne, INRAE, UMR 1095 GDEC, 63000 Clermont-Ferrand, France</footnote>

<footnote n="13">¹³ Carnot Qualiment, INRAE, 44000 Nantes, France</footnote>

<footnote n="14">¹⁴ INRAE, Centre Pays-de-la Loire, Dept TRANSFORM, 3 impasse Yvette Cauchois, CS 71627, 44316, Nantes Cedex 3 Nantes, France</footnote>

<footnote n="15">¹⁵ Present Address: D-Lab Industry, Saint-Bonnet-de-Rochefort, France</footnote>
**Keywords** Plant biology · Agronomy · Human nutrition · Food · Processing · Socioeconomics · Legume · Cereal · Oilseed · Algae · Protein source · Nitrogen fixation

## Contents

1. Introduction

2. Analysis of the current economic context of plant-based protein production

2.1. The rising influence of plant proteins in Agribusiness: market dynamics, sustainability challenges, and innovations

2.2. Challenges in breeding programs to stimulate genetic innovation and progress for protein-rich diversification crops

2.2.1. The role of joint public-private validation in the registration of new varieties

2.2.2 A specific IP system encouraging the reuse of new traits

2.2.3 A genetic trade-off between desired traits: quality vs productivity

3. Protein production: harnessing the different levels of diversity for a multiplicity of uses

3.1. Diversity of the amino acid composition of proteins in plants

3.2. Protein diversity and quality at the species level

3.2.1 Seed protein content

3.2.2. Protein or amino acid composition

3.2.3. Taste and health concerns

3.2.4. Oilseed meal, a potential resource with specific research needs

3.3. Impact of genotypes (G), environments (E), cultural practices (P) and interactions (G×E×P)

3.4. Crop associations: in spite of many advantages, disruptive cropping practices slow their adoption

4. From proteins to foods: designing food products adapted for diverse consumer needs and demands

4.1. Optimization of protein digestibility for human consumption

4.2. Attractiveness of plant proteins for market development

4.2.1. Modifying processing to improve pulses consumed traditionally

4.2.2. Developing new foods through traditional processes (e.g., fermentation)

4.2.3. Adopting "exotic" products

4.2.4. Adapting processes for mixing pulses and cereals in traditional foods

4.2.5. Using new technologies (e.g., extrusion) to develop new foods

4.3. Algae and Microalgae, a niche with added value

5. General discussion and identification of research priorities

5.1 An urgent need of connecting various disciplines and points of view

5.2 Interdisciplinary research needs

6. Conclusion

Declarations

References

## 1 Introduction

While the world's population was around 7.9 billion people in 2022, demographic growth projections suggest that it will 9.7 billion in 2050. Agricultural production will have to increase, while cultivable land tends to stagnate (FAO 2017). At the same time, food systems are known to be one of the main causes of global change (climate change, changes in land use, depletion of freshwater resources, and pollution of ecosystems) as shown by Springmann et al. (2018) who forecast a 50-90% increase of negative environmental effects of food systems before 2050, going beyond the planetary boundaries. Food systems have to evolve (Berners-Lee et al 2018) since current production systems are being questioned, for example, regarding the negative externalities of livestock breeding or the synthetic inputs applied to increase crop yields. Dietary changes towards healthier, more plant-based diets could help to reduce the negative environmental effects of the food system (Springmann et al 2018). Globally, a transition must occur to reach a sustainable and healthy diet. A sustainable diet was defined by FAO (Food and Agriculture Organization of the United Nations) in 2012 (FAO 2012). All the scenarios pointed to the necessity of changes, for instance, in the TYPHA project (Poux & Aubert 2018) or in the EAT-Lancet Commission (Willett et al 2019). Specifically for the populations of Western societies, who expect both a healthy diet and a preserved environment, the interconnections between health and sustainability should be highlighted (Axelos et al 2020; Caron et al 2018). Among these changes, protein consumption is specifically discussed, and there is a general consensus on the need to reduce animal proteins and increase the proportion of plant proteins (Chen et al 2022; de Boer & Aiking 2019; Poux & Aubert 2018; Willett et al 2019). To increase plant proteins in the diet, sustainable and reliable sources of these nutrients must be identified and developed on a sufficient scale to preserve food security. Agricultural production, while allowing farmers to make a decent living from their activity, must mitigate its environmental impacts with decreased use of carbon-intensive inputs. It must also adapt to climate change

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that multiplies extreme weather events leading to changes in the physical environment and pest and pathogen populations (United Nations 2015). The purpose of this review is to address the systemic changes that need to be made to plant production and food processing systems to increase the share of plant-based proteins in the diet of Europe and North America populations. Because the vegetalization of protein sources in human diets is urgently needed, such a synthesis, bringing together multiple disciplines from economics to nutrition, is required to point out the barriers in each field of application and the potential to bypass them by mobilizing solutions in other areas.

Indeed, faced with these challenges, rethinking our production and utilization of plant protein sources is essential. Proteins are primary macronutrients in the human diet. They are composed of 20 amino acids, nine of which are not synthesized by our body and must be provided by the diet; they are called indispensable. The proteins provided by the diet are necessary for the anabolism of proteins in the body. They also participate with carbohydrates and lipids in the energy supply (Passmore et al 1974). The renewal of proteins in the human organism maintains its structural (parenchymal tissues, muscles, bones, skin, appendages, and others), enzymatic, transport, hormonal, and immune functions, among others. International organizations such as EFSA (European Food Safety Authority) (EFSA 2012) and FAO (Passmore et al 1974; FAO 2012, 2013) recommend an intake of 0.83 g of protein per kilogram per day for an adult and between 10 and 20% of protein in the daily diet. According to a French national study of food consumption (ANSES 2017), the male adult population is at 144% of this target and the female at 127%. Less than 33% of these proteins are of plant origin; most of these come from cereals (bread, pasta, rice), while the share from Fabaceae (pulses, soybean) is less than 1% (Chardigny and Walrand 2016; ANSES 2017; Kairo and Zeghnoun 2019). When analyzing the packaged food supply worldwide with legume ingredients, more than half are based on soya, with a low diversity of pulses used (Salord et al. 2022, 2024). The challenges are as much about changing the demand, requiring the mobilization of multiple levers such as educational and awareness actions, regulation, and taxation, as they are about transforming supply and pursuing the development of scientific knowledge. For instance, over the last decades, more than 60% of the scientific literature on food sciences and technology concerning legumes is specialized on soya (Magrini et al 2019), which does not help in creating sustainable pathways for more species diversity in diets. The plant-based protein-rich analogs of dairy or meat, which have been proposed recently, notably by startups and now more and more by large food companies (Good Food Institute 2023; Vincent & Prouteau 2024), have met difficulties convincing the general public beyond the initial curiosity factor (Eastlake 2024; Varela et al 2022; Vincent

& Prouteau 2024), as evidenced in the USA where the market developed earlier but appears to have reached saturation (Bambridge-Sutton 2024; Fassler 2024). Indeed, addressing jointly taste, convenience, and cost requires mobilizing multiple levers. One of these levers is fermentation as it can decrease anti-nutritional factors and modify taste with limited energy expenditure (Tangyu et al 2019), for example, using the power of microbial consortia to design new products, such as illustrated in Figure 1. Other levers include culinary approaches to extend the range of convenient pulse products (Melendrez-Ruiz et al 2019a), changing food offers in foodservice to change consumer behavior (Magrini et al 2021), or informing consumers on pulse benefits (Marette and Roosen 2021).

Although in absolute terms, plant protein production in the European Union (EU), and more specifically in France, would be sufficient for its population (Table 1), it is strongly biased towards cereals, and therefore, it is not adequate when taking into account other qualitative and systemic factors (European Parliament, 2023), from breeding to agronomic systems. To put these data in perspective, we can consider a European population (27 member states) of 494 million people with an average body mass of 70 kg requiring 0.83 g of protein per day per kilogram, presenting a requirement of 10.5 Mt of protein per year.

EU and France continue to be highly dependent on imported plant proteins because (i) they have a deficit in protein-rich materials (PRM) for animal feed, which implies imports of soybeans and sunflowers in the form of seeds and high-protein oil meals, (ii) they struggle to meet the needs of dry peas or faba beans for functional uses in human food, and (iii) the need for pulses (lentils, chickpeas, beans) is not met while being below recommended consumption levels. To face this situation, the French government launched a new plan in 2021 (Ministère de l'Agriculture et de la Southernaineté Alimentaire 2021) of nearly 100 million euros of public aid to support the production of plant proteins and strengthen protein sovereignty.

Increasing the production and consumption of plant protein sources can contribute to a healthy and sustainable human diet (Willett et al. 2019) by providing in particular indispensable amino acids, fiber (Hertzler et al. 2020), vitamins, and minerals (Mudryj et al. 2014) as well as phytochemical compounds. It will also accompany a decrease in consumption of foods from animal origin, whose excess is associated with an increased risk of chronic diseases (Godfray et al. 2018). Moreover, locally produced plant proteins could contribute to developing new economic activities. From an environmental point of view, Fabaceae offer ecosystem services in single crops or intercropping systems thanks to their capacity to fix atmospheric nitrogen, which enriches soils with nitrogenous compounds. Pulses and other legumes are still struggling to expand in the human

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**Fig. 1** “Camenverts”, an example of dairy alternative products prepared with pea protein isolates (Saint-Eve et al 2021) and endowed with sensory properties (texture, aroma) close to those of typical French cheeses by fermentation using selected microbial consortia. Photo credit: Françoise Irlinger

<figure><img src="image_6.png" /><figcaption>**Table 1** Protein resources in the EU-27 (average 2016–2020). Sources: Eurostat for EU-27 production (Eurostat 2022) and Feedipedia for protein content (Feedipedia 2012); calculations from the authors.</figcaption></figure>

<table><thead><tr><th>Source</th><th>Annual harvest (Mt)</th><th>Protein %</th><th>Protein weight (Mt)</th></tr></thead><tbody><tr><td>Wheat and spelt</td><td>131.60</td><td>10.9%</td><td>14.34</td></tr><tr><td>Corn grains and corn-cob mix</td><td>66.87</td><td>8.1%</td><td>5.42</td></tr><tr><td>Barley</td><td>53.12</td><td>10.0%</td><td>5.31</td></tr><tr><td>Rapeseed and turnip seeds</td><td>17.55</td><td>19.0%</td><td>3.34</td></tr><tr><td>Triticale</td><td>11.26</td><td>9.9%</td><td>1.11</td></tr><tr><td>Oats and mixtures</td><td>10.14</td><td>9.2%</td><td>0.94</td></tr><tr><td>Sunflower seeds</td><td>9.72</td><td>12.7%</td><td>1.23</td></tr><tr><td>Rye and winter cereal mix</td><td>7.98</td><td>8.8%</td><td>0.71</td></tr><tr><td>Rice</td><td>2.89</td><td>7.9%</td><td>0.23</td></tr><tr><td>Soybean</td><td>2.74</td><td>35.5%</td><td>0.97</td></tr><tr><td>Dried peas</td><td>2.17</td><td>20.4%</td><td>0.44</td></tr><tr><td>Broad beans and fava beans</td><td>1.18</td><td>25.1%</td><td>0.30</td></tr><tr><td>Sorghum</td><td>0.88</td><td>9.2%</td><td>0.08</td></tr><tr><td>Sweet lupins</td><td>0.22</td><td>33.5%</td><td>0.07</td></tr><tr><td>Total</td><td>318.33</td><td></td><td>34.49</td></tr></tbody></table>

diet due to complex preparation and a lack of sensory appeal (Melendrez-Ruiz et al. 2019a, b). In particular, they are still strongly constrained by sensory off-notes. Niche markets with varieties adapted to their terroirs would make it possible to develop territorialized food systems while being able to contribute to supplying agro-industrial chains intended for

export. Another possibility could be to increase the recourse to algal proteins, as macroalgae do not compete for land use. However, this has met with limited interest so far: while macroalgae are currently harvested or grown for functional polysaccharides (agar, alginates, carageenans), recent developments have focused more on microalgae for other nutrients (polyunsaturated lipids, carotenoids). The diversity of markets associated with the demand for plant proteins constitutes multiple opportunities for the joint restructuring of agricultural and food systems.

The multiple challenges faced to enable this multifaceted transition, and their interconnections, are depicted from farm to fork in Figure 2. The first and second blocks relate to protein production, the first including genetic resources and agronomical conditions and the second the characteristics of the resulting products. The third and fourth blocks deal with proteins in food products, the third block being the processing of the diverse sources of plant proteins, with a priority but not exclusivity on pulses, while the fourth depicts what is known on the characteristics of these products from a consumer point of view. These four blocks include different disciplines in which research is carried out about plant proteins. The current paper will focus specifically on the research topics for the use of plant proteins in food.

So far, very limited inter- or multi-disciplinary research has been carried out on plant proteins. However, as depicted in Fig. 2, interconnections and retroactions need to be understood to inform public policies for transitioning to a plant-based diet in Western societies. Because operational solutions cannot be found by

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**Fig. 2** Schematic representation of the research required, from production to consumption, from farm to fork. On the left are the different levels of crop diversity; the crosses symbolize the possible interactions between these levels. The yield and composition of plant production, the suitability for the various processes, and the multiplicity of uses are criteria to be considered. The choices of compositions, processes, and uses will have feedback, symbolized by the arrows, on the cultivated diversity, at the scales of the territories and the plots

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addressing only one step in this complex web, scientists from the public and private sectors decided to write this synthesis, which reviews recent advances, identifies collective research priorities, and contributes to coordinating this research upstream and downstream of plant protein production chains.

# 2 Analysis of the current economic context of plant-based protein production

## 2.1 The rising influence of plant proteins in agribusiness: market dynamics, sustainability challenges, and innovations

The constant increase in cumulative investments in these booming markets proves the growing enthusiasm of agribusinesses for the plant protein market. The consumption of plant proteins is growing (FAO 2022), as is their market (ALCIMED 2022, Morach et al. 2021), and should continue to do so until 2035 (ALCIMED 2022; Eastlake 2024; Morach et al. 2021). The world market for plant proteins represented 7 billion euros in 2013, 10 billion euros in 2018, and will reach 15 billion euros in 2025 (ALCIMED 2022). This trend can be explained by the awareness of the need to increase plant-based protein consumption to substitute part of animal proteins. However, this production of plant proteins should be based on diversified agroecosystems with low use of agrochemical inputs to be sustainable (Berners-Lee et al. 2018; FAO 2012; Springmann et al. 2018). While there is currently a strong development of protein isolates from major world crops (gluten from wheat, tofu and its derivatives from soybean, Salord et al. 2022), more is needed for minor diversification crops such as pulses.

Among these diversification crops, pea protein remains the most used protein for food after wheat and soybean in the world and in Europe (Salord et al. 2022). Pea proteins can be extracted using technologies adapted from those initially used for pea starch production (Dijkstra et al. 2003), such as milling or suspension in an alkaline solution and centrifugation, followed by pH-shift for protein precipitation. This process could also be deployed for other crops. For example, in France, DSM and AVRIL companies (Haverland 2021) launched Olatein, the first industrial plant producing rapeseed-based protein based on a technology that avoids using solvents during the defatting stage.

Pulses compete with the major cash crops such as wheat and soybean, which are still the ones from which most plant proteins for food innovations are extracted, as evidenced by research (Chardigny and Walrand 2016; Guéguen et al. 2016; Lam et al. 2018) and market studies (Bashi et al 2019; EUVEPRO 2019; Rabobank 2017; Salord et al. 2022). In 2017, only 2% of the ingredients used to replace meat in the diet came from grain legumes other than soy (Magrini et al. 2023).

The development of biodiversity cultivated for plant proteins is a crucial issue questioning the capacity of the market to overcome path dependency effects leading to the lock-in of this system (Conti et al. 2021; Meynard et al. 2018). Path dependency refers to the fact that decisions taken in the past can generate self-reinforcing dynamics, with a high cost of change at a given moment preventing profitable evolutions in the long run. Here, it refers specifically to the priority in cereal self-sufficiency following the Second World War, as supported by the Common Agricultural Policy, and the international negotiations of the Uruguay Round (Abis 2023). The performance of the initial production choices, reinforced over time through the effect of self-reinforcing mechanisms (known as “increasing returns
to adoption”), has widened the competitive gap between major and minor crops by focusing genetic and agronomic research and development on the major crops (Magrini et al. 2016). Today, this gap can no longer be reversed by market forces alone. Private breeders are reluctant to invest in orphan species because this is a narrow and risky market with little investment from the downstream sectors. Producers invest little in them because the upstream production is unstable and underdeveloped. Therefore, reversing such lock-in calls for specific investments in minor crops so that all operators gain an economic interest in their development (Association Nationale des Industries Alimentaires 2021; Magrini et al. 2018; Cusworth et al. 2021a). The diversity of seed legume species is vast, leading to a large repartition of research investments between winter and spring peas, winter and spring faba beans, white lupin, narrow-leaf lupin (blue), yellow lupin, chickpeas, lentils, dry beans, and, of course, soybeans with all their precocity ranges, so as to find an appropriate niche in crop rotation. An essential step in developing species not widely cultivated today would be to increase the selection effort by devoting specific resources from the public sector. Thus, public regulation should support investments along value chains to consolidate the competitiveness of these minor species, which have received fewer investments until now.

If market innovations are numerous, opening up the field of possibilities also implies that companies first have access to strategic information such as market one. Most market studies are carried out by and for large agro-industrial groups, but SMEs get little access to such analyses to think about the future while they constitute 98% of the agri-food sector in France (Marette and Roosen 2021). Faced with this asymmetry in information, the role of public and interprofessional organizations is crucial in sharing market knowledge. This knowledge is based mainly on recent research that analyzed market biodiversity through the diversity of plant species used in food product launches (Salord et al. 2022, 2024).

The impact of consumer information about the environmental benefits of plant proteins on purchase forecasts should also be considered. However, recent studies show that this environmental information has little effect on modifying consumption habits towards legumes (Melendrez-Ruiz et al. 2019a) and that sensory determinants and culinary culture (Alston et al. 2012; Melendrez-Ruiz et al. 2019b) remain major. Emphasizing proven or potential health effects, with the perception of an individual benefit, could also be a way to influence these choices. Furthermore, improving the properties of plant proteins is necessary for them to better meet acceptability criteria (Alston et al. 2012; Melendrez-Ruiz et al. 2019b). This requires transdisciplinary research

to identify targets for improvement and methods to either increase the content or bioavailability of nutritionally desirable nutrients or decrease the abundance of nutritionally undesirable molecules. This can be achieved either through genetics/breeding or processing/cooking, or combining both approaches, depending on the use of seeds, flour, or ingredients and the role of such molecules in plant defense against biotic or abiotic stresses.

## 2.2 Challenges in breeding programs to stimulate genetic innovation and progress for protein-rich diversification crops

Meeting society's needs in the context of agroecological and food transitions and climate change requires substantial investments in research and innovation. In the French plant breeding sector, the innovation ecosystem is structured around three complementary and interdependent sets:

i. Registration in the Official Catalogue constitutes authorization to market new varieties. This tool guarantees the adequacy of the varietal offer to the needs of farmers and sectors while serving as a lever for the orientation and timing of genetic progress (see Section 2.2.1).

ii. The intellectual property (IP) protection system (see Section 2.2.2) for new varieties based on the Plant Variety Certificate (PVC) gives companies the means to invest in R&D work to meet the needs and challenges according to the registration guidelines.

iii. Finally, varietal innovation is based on exploiting genetic diversity, maintained in biological resource centers or in situ, and constitutes the raw material for genetics and breeding work.

### 2.2.1 The role of joint public-private validation in registration of new varieties

In France, the access to the market of a new variety relies on its registration in the French Official Catalogue and then, by extension, in the European Catalogue. This registration depends on regulations established at the European level, whose implementation in France is entrusted to a joint public-private, participatory body, the Comité Technique Permanent de la Sélection (more details on https://www.geves.fr/qui-sommes-nous/ctps/). It plays a significant role in implementing and evolving the registration procedures to obtain adequacy between the objectives of the various stakeholders, i.e., farmers, users, companies and public authorities, and the scientific and technical capacities of the breeders to meet the needs of agriculture and its sectors.

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To be registered, a new variety must satisfy the so-called DUS tests: Distinctness, Uniformity, and Stability. That is to say that a registered variety must be distinct from the varieties already marketed and their characteristics must be stable through multiplication processes and homogeneous. Moreover, for some species, it must also be an improvement compared to the existing one. This improvement is what the Value for Sustainable Crop Use (VSCU) examination seeks to officially qualify, based on field trials and technological tests, jointly assessing agronomic criteria (including yield, resistance to pests and diseases, nutrient, and water use efficiency, among others) and the quality of harvested products (protein content, oil content, forage quality, among others). These criteria are examined through a network of official trials implemented by the national testing office, GEVES (more details on www.geves.fr). The evaluation criteria and the level of performance that new varieties must achieve compared to existing varieties allow VSCU to guide, set the pace, and frame genetic progress.

### 2.2.2 A specific IP system encouraging the re-use of new traits

To allow companies that invest in plant breeding R&D to earn a return on their research efforts and to encourage these investments, an IP protection system has been built and implemented specifically for plant varieties under the International Union for the Protection of New Varieties of Plants (UPOV). It allows the breeder to claim ownership of a variety as an original set of traits through DUS and novelty testing. In a *sui generis* IP protection system, the Community Plant Variety Right (CPVR) is a lever for promoting innovation. It allows companies to be remunerated for their R&D efforts by giving the entire innovation ecosystem access to new varieties (breeder's exemption). Indeed, a new protected variety can be used by any other breeder as a genitor in his breeding programs without the initial creator being able to claim a right to the varieties that come from these genitors. This system is a strong incentive to invest in R&D for companies devoting 13-15% of their turnover to innovation and breeding.

Thus, the innovation ecosystem built around varietal innovation makes it possible to both qualify and direct genetic progress through registration while at the same time giving breeders the means to innovate to meet the registration criteria (e.g., the positive Grain Protein Deviation GPD+ criterion, see Section 2.2.3 below). This virtuous circle is nevertheless confronted with different limits for which solutions are sometimes found. In the French system, the CPVR exemption system allows farmers to use part of their own harvest from a protected variety to reseed their fields

in return for a fair and equitable payment to the breeder of the variety. This practice of farm-saved seeds is widespread in self-pollinating crops such as wheat and peas. The implementation of this system in France, provided by Article 14 of Regulation 2100/94/EC (Council Regulation 1994), has been established for straw cereals (bread wheat, durum wheat, barley, triticale, oats, spelt, rye, and rice) through an interprofessional agreement, renewed in 2019, which sets the terms and amount of a Contribution to Research and Varietal Innovation (CRIV) levied on the collection of straw cereals. The breeders have jointly decided to invest 15% of the total amount collected for their benefit in public-private research projects for the genetic progress of all straw cereal species. These projects are implemented within the fund's framework to support plant breeding. This fund is a virtuous approach to pooling private funds to support pre-competitive research projects in collaboration with public research, but it only concerns straw cereals.

Its implementation for grain legume species, also self-pollinating, is confronted with (i) the great diversity of these species, (ii) their small area of cultivation, and (iii) the limit of its implementation cost, which becomes too high given the small volume of harvests. Like other mechanisms implemented, this situation leads to a strengthening of R&D investment in improving specific species. It accentuates the gap with minor crop species, whose competitiveness and relative attractiveness for private breeders and producers are decreasing, feeding a logic of collective disengagement.

In the context of market failure, public intervention is justified to correct the imbalances observed. Thus, the breeding of varieties for minor crops has traditionally relied on public research and innovation forces, notably implemented within INRAE through the "Variatetal Innovation and Diversification" programs in partnership with its subsidiary Agri Obtentions. However, the resources invested remain below the stakes: between 2014 and 2024, 333 varieties of bread winter wheat were added to the Catalogue, but only 66 varieties of winter pea (see https://www.geves.fr/catalogue/ for data, calculation by the authors). It would be possible to imagine specific support dedicated to public varietal creation programs from the European Union within the framework of the Farm to Fork strategy of the Green Deal. The diversification of cultivated species is indeed a challenge for food security, pesticide-free agriculture, and adaptation to climate change on a European scale. Further, this support should be provided over a period compatible with the creation of varieties, which is 7 to 15 years. This is far from the traditional 3 years frame of calls for projects. This also implies innovative organization and funding structures to enable the socio-technical unlocking (Paux et al. 2022) of the disability experienced by minor species.
Over the past decade, France has made a significant con-
tribution to the support of plant breeding through ambitious
research projects launched within the framework of the
“Programmes d’Investissement d’Avenir” (PIA—Investment
Programs for the Future), such as Breedwheat for wheat,
Amaizing for maize, Peamust for protein peas, Sunrise for
sunflower, or Phenome for the development of phenotyp-
ing infrastructures. Based on a public-private partnership to
optimize agreed-upon resources, these programs have led to
significant advances for major crops, as shown for wheat and
reviewed by Paux et al. (2022). Nevertheless, many of the
results of the work carried out within the framework of the
PIAs still have to be translated into transfers and innovations
for companies and sectors. The challenge facing the plant
breeding ecosystem today is to translate the results of this
high-quality research into innovations, markets, and solu-
tions for the benefit of farmers and value chains.

2.2.3 A genetic trade-off between desired traits: quality vs productivity

A further challenge is that some of the aims of breeding
innovation may not be compatible, as highlighted here in the
example of productivity and the protein content of wheat. For
wheat, given the VSCU tests in France, any new variety must
meet criteria for grain yield, pest resistance, protein concen-
tration, and technological quality. However, the joint improve-
ment of yield and protein concentration is difficult due to the
strong negative correlation between these traits (Figure 3):

the more a variety is productive, the more diluted are proteins
in its grains, and the lower is the protein content. To counter
this effect, breeders are working on the GPD (Grain Protein
Deviation) whose genetic basis has been analyzed (Bogard
et al. 2010; Taulemesse et al. 2015) (see Section 3.2.1).

The GPD is a good example of the interaction between
regulation and breeding priorities. The GPD is considered
by the CTPS thanks to a bonus/malus system modulating
the final score at registration and encouraging the selection
of productive varieties with a protein concentration higher
than the one predicted by the regression (GPD+ varieties).
Through this bonus system, the CTPS can orientate breed-
ing activities and genetic progress so that the new varieties
meet the market's expectations, which are strongly seg-
mented today. All segments of the wheat market (except
cookie wheat) require a high protein concentration and
are, therefore, very dependent on fertilizers to ensure non-
limiting nitrogen nutrition. The bonus system has worked
well, and today's French varieties meet the dual challenge
of productivity and protein concentration. However, these
efforts to overcome the negative correlation between grain
yield and protein content would require better payment for
the production of protein-rich wheat. GPD is widely used
by geneticists and by farm advisors. It can contribute to the
improvement of the economic model by helping to adjust
the varietal choice to the market segments, in the differ-
ent territories (feed wheat, bread wheat, durum wheat,
export, short circuits, among others), in an agroecologi-
cal perspective, i.e., without systematically looking for a

<figure><img src="image_10.png" /><figcaption>**Fig. 3** Schematic representa-
tion of the negative relationship
between yield and protein con-
centration, used to estimate the
Grain Protein Deviation (GPD)
of varieties in wheat, from
Taulemesse et al. (2015). Each
diamond represents a variety.
The blue, green, and red: varie-
ties behave as expected, with
a protein concentration higher
(GPD+) or lower (GPD-) than
expected from the regression.
MS: dry matter content</figcaption></figure>

<figure><img src="image_11.png" /></figure>
high protein content at the cost of a strong dependence on nitrogen fertilizers.

A concluding remark on this chapter about plant genetics is that few quality criteria concerning human food are currently explicitly considered for variety registration (GPD, as presented above, some anti-nutritional factors or mycotoxins...). Integrating such standards could be another key in unlocking the potential of pulses for diets.

## 3 Protein production: harnessing the different levels of diversity for a multiplicity of uses

The effects of diversification appear generally positive at many levels and have been studied and reviewed from the experimental plot (Beillouin et al. 2021) or the farm (Furman et al. 2021) to whole countries (Renard and Tilman 2019) and through a recent collective scientific expertise (Vialatte et al. 2021). Diversity of crops is highly relevant to sustainability as stability of harvest in a given context is strongly linked to diversity of cultivated species, as has been shown on a large historical and geographic database (Renard and Tilman 2019). Diversity can also be harnessed to ensure nutritional balance, as is done for proteins by combining legumes and cereals (see Section 3.2.3). There are however discussions on whether to promote diversification at the global level, linked to economies of scale and processing, or diversification at the local level.

### 3.1 Diversity of the amino acid composition of proteins in plants

The quality of protein for human nutrition can be characterized by its amino acid composition. Different amino acid profiles are recommended for different ages (WHO 2007). Protein quality also depends on the bioavailability of the amino acids, particularly protein digestibility. When digestibility corrects the relative amino acid composition, it is possible to determine the indispensable amino acids available for protein synthesis after their intestinal absorption (Ennis and Elango 2021; Reeds 2000). The DIAA ratio (digestible indispensable amino acids) indicates whether each indispensable amino acid's level is higher or lower than the requirements. The lowest ratio among the nine indispensable amino acids defines the DIAAS (DIAA score) (FAO 2013). The DIAAS of plant proteins is often < 1 (Herreman et al. 2020) because most have at least one limiting amino acid. Lysine is the most limiting amino acid among plant proteins from cereals but also from nuts or some oilseeds. Legumes are generally less deficient, with methionine being the most limiting amino

acid, but other amino acids are sometimes present in small amounts, such as valine, leucine, and isoleucine for faba beans. Some plant proteins show little or no deficiency. The latter situation is valid for rapeseed, potato, quinoa, pea, and soy proteins (Herreman et al. 2020). In recent decades, supplementation between multiple incomplete protein sources has become a paradigm. An advantage of DIAAS is that it allows the calculation of the DIAA ratio for a combination of ingredients in a food or meal. The additivity of DIAAS has been demonstrated experimentally (Fanelli et al. 2021). Herreman et al. (2020) virtually optimized DIAAS with a combination of plants, for example, a potato ration (65%) with corn (20%) and faba bean (15%) to achieve 100% DIAAS. This principle can be applied to various meal combinations to ensure that diets provide sufficient digestible amino acids consistent with the crop production area.

Protein bioavailability varies according to the food matrix (e.g., protein isolates being more digestible than seeds) and the recommended amino acid profile (e.g., children between six months and three years old). The seed amino acid profile can also be influenced by nutrient deficiencies and environmental factors, as described in Section 3.3.

While the amino acid composition profile is a critical factor in nutritional balance, most plant proteins are not consumed pure but in processed foods that affect their bioavailability and determine their attractiveness to the consumer. Factors such as taste, texture, ease of use, color, and the presence of flatulence-producing galactosides are far more critical in determining purchasing behavior than nutritional balance (Roland et al. 2017). Valuation in short chains by a network of local SMEs will probably be the best way to promote varietal improvements and develop market segmentation (i.e., partitioning the customers into smaller groups that share similar characteristics, such as age, income, personality traits, behavior, interests, needs, or location, and targeting their needs in a specific rather than general fashion) to create value by carrying niche innovations from one end of the chain to the other. For instance, this was done successfully in France with green lentil "Lentille du Puy" under a Protected Designation of Origin, unfortunately restricted to a single variety (for more information http://www.lalen.tillevertedupuy.com/). This approach is complementary and competitive with producing more purified, processed ingredients such as isolates and concentrates.

### 3.2 Protein diversity and quality at the species level

#### 3.2.1 Seed protein content

Seed proteins determine the amino acid balance. Thomas Burr Osborne (1859–1929) laid the foundations for the biochemistry of proteins, defining a classification based on their
solubility in different solvents, which is the basis for the extraction processes still used today. Four protein classes can be distinguished: albumins soluble in water, globulins soluble in a saline environment, prolamins soluble in an alcoholic solution, and glutelins soluble in alkaline conditions. Each of these protein groups is further subdivided into several protein fractions, which differ in their structure and amino acid composition. Seed storage proteins (SSP) are mainly stored in the reserve tissues, either the embryo (in the case of legume seeds) or the endosperm (in the case of cereals).

For cereals, the major SSP are prolamins and glutelins (Lam et al. 2018). The term “prolamins” comes from the amino acid composition of this protein class, which was reported to be generally rich in proline and amide nitrogen derived from glutamine in most cereals with the exception of rice and oat. The combined proportions of these amino acids range from about 30–70% of the total among different cereals and protein groups (Shewry and Halford 2002). In wheat, prolamins and glutelins are the gliadins and glutens, respectively. They are also called gluten proteins because they aggregate after hydration and kneading to form the gluten network that gives dough its technological qualities for processing. Indeed, a large part of the wheat production is consumed by humans after processing into bread, noodles, or pasta (for durum wheat). The current industrial versions of these processes require a high protein content and an adapted gliadin to glutenin ratio (Shewry 2024).

In legumes, globulins are the most abundant SSP and are classified into the 7S and 11S globulins. The ability of pulses to offer a high protein content in the absence of nitrogen fertilization makes them particularly attractive as a source of plant protein, contributing simultaneously to the agroecological and nutritional transitions. Soybean-based products that go through the formation of colloidal suspensions (tonyu, “juice” obtained by grinding soy beans in water and centrifuging) represent the main soybean food uses in France: they are the source of consumption of 45,000 tons of seeds per year (Terres Univia 2021). The yields of tonyu/okara (the solid residual part after centrifugation of the preparation of tonyu or soy milk) production depend on the 11S to 7S globulin ratio in addition to the seeds' protein content (Cai and Chang 1999). This 11S/7S ratio is also crucial for formulating tofu (curdled tonyu), as 11S-rich tofu is firmer and has a higher water retention (Saio and Watanabe 1978).

It is important to distinguish between the total protein content and its composition (i.e., the contribution of each protein class), as the two characteristics are not interrelated. Interspecific and intraspecific variations can be exploited to improve these two characteristics. For instance, legume proteins with high protein and lysine content are well complementary to cereals, which are rich in sulfur-containing amino acids (cysteine and methionine) (more detailed

explanation in Section 3.2). Different varieties of the same species can have different SSP contents and compositions. For example, in legumes, there are wide intra- and interspecific variations in seed protein content, ranging from 16 to 32% in peas, 18 to 32% in lentils, 22 to 38% in faba beans, and 26 to 57% in soy beans (Burstin et al. 2011). These wide variations within each species open the possibility of genetically increasing protein content. This situation is expected to be achieved without lowering yield. In pea, the correlation between yield and protein content ranges from -0.47 to 0.33, depending on the environment and genetic background (Burstin et al. 2007; Klein et al. 2020). Regions of the genomes that control protein content have been identified in legumes, such as peas and soybean (Klein et al. 2020; Teng et al. 2017; Warrington et al. 2015). In soybean, parental lines have been identified that are a good source of alleles for improving both seed protein content and yield (Cober and Voldeng 2000), and genes controlling protein content have been identified (Fliege et al. 2022; Zhang et al. 2020). These genes and loci can be used to meet breeding objectives for improving seed protein content.

Improving SSP concentration without reducing yield is difficult in cereals due to the negative correlation between seed yield and protein content (Figure 2). However, recent studies have identified loci in wheat that increase protein concentration without decreasing yield. One of these loci improves the remobilization of nutrients such as nitrogen, iron, and zinc from the leaves to the grain by affecting plant senescence. It carries the *NAMBI* gene, which encodes for a transcription factor regulating senescence (Uauy et al. 2006). Because protein synthesis is highly dependent on nitrogen fertilization, reducing nitrogen demand while maintaining high yields by improving nitrogen use efficiency is a major concern, as reviewed in rice (Hu et al. 2023) and wheat (Cormier et al. 2016), where progress has been made in dissecting the genetic determinism of nitrogen use efficiency as well as in wheat (Bogard et al. 2010; Taulemesse et al. 2015).

### 3.2.2 Protein or amino acid composition

The need to study protein quality from pulses is becoming increasingly clear, especially regarding the requirements of indispensable amino acids in humans. The amount of protein from pulses to be consumed to cover the daily needs of an adult for each indispensable amino acid is given by Leinonen et al. (2019). It suggests that this balance can be achieved in pea by increasing the proportion of sulfur-containing amino acids (S-AA) by 30%. The genetic variability in the amount of S-AA in pulses (Bastianelli et al. 1998; Gabriel et al. 2008) can be exploited to understand the genetic basis for breeding programs, as has been done in soybean (Malle et al. 2020; Warrington et al. 2015). Genes controlling the

<figure><img src="image_12.png" /></figure>
synthesis of SSP richer or less rich in indispensable amino
acids are among the candidate genes to be considered for
improving amino acid balance. In peas, variations in SSP
composition between genotypes (Bourgeois et al 2009;
Gueguen and Barbot 1988; Tzitzikas et al. 2006) have been
exploited to identify regions of the genome that control this
trait (Bourgeois et al. 2011). The model legume *Medicago
truncatula* Gaertn. whose genome shows good synteny with
that of pea (Tayeh et al. 2015) was used to propose can-
didate genes at these loci (Le Signor et al. 2017). Among
them are potential regulators of globulin synthesis (Le
Signor et al. 2017), including the transcription factor ABA
Insensitive 5 (ABI5) that controls 7S globulin synthesis in
beans (Ng and Hall 2008; Pandurangan et al. 2016). In peas,
mutations in the ABI5 gene significantly reduce the level of
vicilin-like 7S globulins (Le Signor et al. 2017), which are
poor in S-AA and less advantageous than 11S globulins for
emulsion formation (Barac et al. 2010). Additional work is
needed to identify the gene networks controlling AA and
protein composition of seeds and to identify a combination
of alleles to improve high-protein pulse varieties.

Cereal proteins are all low in lysine (Hertzler et al. 2020).
Rice, barley, and wheat are rich enough with S-AA, which
is not the case for maize. Genetic research has been carried
out to correct lysine deficiency in cereals, taking advantage
of genetic variability at the species level. For example, the
lysine content in wheat varies from 0.36 to 0.51 mg per
100 g of whole meal flour, which is higher in diploid or
tetraploid wheat (Singh et al. 2001). Studies of lysine-rich
mutants have shown the role of specific transcription fac-
tors involved in SSP synthesis, such as Opaque2 in maize
and PBF in barley and wheat (Moehs et al. 2019). Muta-
tions rendering these transcription factors non-functional
generate genotypes with less SSP and more lysine-rich
proteins (Habben et al. 1995; Singh et al. 2001; Vaag and
Munck 1987). Other transgenic approaches have been used
in cereals and dicotyledons to increase lysine concentration
by targeting:

1. Enzymes of the lysine biosynthetic pathway, for example, the enzyme catalyzing the first step of this pathway. As reviewed by Wang et al. (2018), in higher plants, the rate-limiting enzymes in lysine biosynthesis are the aspartate kinase (AK) and dihydrodipicolinate synthase (DHDPS), which are extremely sensitive to feedback inhibition by lysine. In founder works, Frizzi et al. (2008) increased the lysine concentration of maize grains transformed for the expression of several biosynthetic pathway enzymes by a factor of 40, while a corn event called LY038 overexpressing the DHDPS was developed for food and feed used in several countries. Overexpression of key enzymes of lysine pathway was also successfully used in rice (Yang et al. 2021) leading

to an increase in free lysine level in transgenic lines by
a factor ranging from 39.2 to 58.5.

2. Overexpression of lysine-rich proteins (see examples in Shewry (2007)).

High-throughput genotyping techniques are accelerat-
ing the discovery of genetic markers for use in breeding
programs (Tayeh et al. 2015). Since all the necessary tools
(marker s and reference genome sequence) are available or
are being developed in most cereals and grain legume crops
(Jayakodi et al. 2023; Kreplak et al. 2019; Tayeh et al. 2015;
Xie et al. 2019), genome-wide association studies (GWAS)
can be performed to determine the chromosomal regions,
or even the genes, involved in complex traits, such as amino
acid balance. Such an approach has been used in soybean,
where candidate genes for improving seed methionine con-
tent have been identified (Malle et al. 2020). It has also been
used recently in wheat to determine the genetic control of
free asparagine (Lavoignat et al. 2024), which is positively
associated with the content of acrylamide, a probable car-
cinogen to humans (IARC 1994). However, these studies
require large numbers of individuals (>200) to obtain reli-
able results, so they are generally limited by the phenotyping
capacity of complex traits. In addition, as mutant libraries
are also available, reverse genetics can be applied to validate
gene function.

3.2.3 Taste and health concerns

Wheat SSP trigger coeliac disease (Sapone et al. 2012) and can provoke allergies (respiratory allergy, food allergy like wheat-dependent exercise-induced anaphylaxis, and contact urticaria (Sapone et al. 2012)). While allergies can be also induced by albumins-globulins, the role of gluten in non-coeliac wheat sensitivity is still discussed (Sapone et al. 2012). However, specific wheat proteins involved in adverse immune responses and clinical manifestations are well described and have been mapped on the wheat genome (Juhász et al. 2018).

In soybean, the presence of isoflavones, which have phy-
toestrogenic properties, makes them endocrine disruptors,
an industry concern. A 2005 AFSSA report (AFSSA 2005)
recommends a dialy intake limit for isoflavones of 1 mg/day/
kg, which has been reached by some consumers who have
replaced dairy products with the family of soy derivatives
(Que Choisir 2019). As isoflavones play a role in nodulation
(Cleyet-Marel et al. 1996), it remains possible that reducing
their content in the seeds could induce adverse effects on the
vegetative vigor of the plant. Anti-nutritional factors such
as lectins and antitrypsin factors seem to be of little concern
for soybean products.

A survey carried out by Terres Inovia among the fore-
most industrialists of this sector has highlighted a certain
number of expectations regarding seed quality. Organoleptic acceptability is a critical factor. Each batch can only be used if it passes the sensory test. This quality is inseparable from a manufacturing process, the negative flavors resulting mainly from oxidative fat degradation in the seed by lipoxygenase (LOX), a key enzyme in the oxidative degradation pathway of lipids. This LOX pathway leads to aldehydes, such as hexanal, responsible for “herbaceous” or “green” odors (Baysal and Demirdoven 2007; Robinson et al. 1995). It is traditionally inactivated by heating, but these treatments have the undesirable effect of reducing the solubility of proteins (Adams 1991; Borhan and Snyder 1979; Liu 1997). Fully acetylated group A saponins, located generally at the hypocotyl, are the primary cause of undesirable bitterness and astringency in soybean foods. The degree of acetylation affects the intensity of the undesirable characteristics of these products, such as bitter taste, astringency, and green bean flavor (Chitisankul et al. 2018; Okubo et al. 1992; Yoshiki et al. 1998).

Breeding in legumes has been aimed at removing certain anti-nutritional or harmful compounds from the seeds, like trypsin inhibitors in peas (Clemente et al. 2015) and tannins or vicin/convicin in faba beans (Duc et al. 1999; Ray et al. 2015). The biosynthesis of saponins that contributes to the bitterness of proteins extracted from pulse seeds has recently been inhibited by targeting the first enzyme of this pathway by TILLING (Targeted Induced Local Lesions IN Genome) in pea (Vernoud et al. 2021), thus providing new alleles for varietal improvement.

### 3.2.4 Oilseed meal, a potential resource with specific research needs

Oilseed meals (rapeseed, soybean, and sunflower), usually intended for animal feed, represent a potential source of concentrated proteins for human food. These meals could be used to manufacture protein-rich ingredients to accompany vegetables and cereals to balance the amino acids intake. At present, few uses of this type have been identified. Soy proteins textured by extrusion after mechanical defatting are available on the market and are incorporated into wet preparations to provide a texture close to that of minced meat (Guyony et al. 2022). The disadvantage of these products is that they remain rich in oxidizable lipids, isoflavones, and oligosaccharides. Further processing would be required to reduce the concentrations of these compounds. Oilseed meals produced as by-products of oil production are of insufficient quality for use in human food. Several factors are involved in this unsuitability.

The first is the thermal degradation of proteins. This process occurs in the meals' desolvation stage, in which the product is exposed to temperatures of ~105 °C for tens of minutes (Carré 2021). This results in non-enzymatic

browning, loss of part of the lysine in rapeseed (15-20%), reduced protein digestibility due to complexes with phytates and fibers, and the formation of unattractive toasted flavors (Salazar-Villanea et al. 2016).

Another undesirable characteristic is the presence of the lignin-rich envelopes of the seeds (rape and sunflower), which produce an inedible product with an unpleasant texture.

Rapeseed also contains several undesirable substances responsible for astringency (tannins), bitterness (glucosinolates derivatives, sinapine, and polyphenols), foul odor, and anti-nutritional activity (goitrin). Consequently, rape-seed protein requires a high level of purification to be used without problems in human nutrition and does not seem to be a target for simple processing (Carré 2021). However, an industrial plant for producing rapeseed proteins for food has opened in Dieppe (Haverland 2021; Olatein 2022). In terms of genetic improvement, there is no simple route to significantly improve the prospects in this area, as there are so many traits to introduce. A breeding program supported by the French oilseeds organization focuses on improving the protein content of winter rapeseed at the expense of fiber and lignin content (Terres Univia 2020). On the other hand, glucosinolate contents have stagnated for more than a decade. Breeders find improving traits related to productivity and resistance to plant pests and diseases challenging while reducing seed glucosinolates.

In contrast, sunflower has no anti-nutritional factors apart from a certain amount of phytate. However, two points limit its interest:

- It would be necessary to shell the achenes entirely and remove as much of the oil as possible by mechanical extraction. However, the first step is impossible to achieve with correct production yields. Increased oil content in commercial hybrids was achieved by modifying the achene composition, favoring a reduction in the proportion of the pericarp to that of the embryo (Denis et al. 1994). A side effect is that these achenes are much more resistant to hulling, mainly because of the adhesion phenomena between the hull and kernel. Significant variations in the hulling ability within cultivars (Dauguet et al. 2016) could be exploited to limit adhesion.

- The sunflower turns dark during crushing due to the oxidation of chlorogenic acid (Sastry and Rao 1990). However, this natural phenomenon does not a priori have undesirable effects on nutritional quality.

Commercially, mechanical defatting can reduce the residual oil content to less than 7%, and the meal can contain more than 50% protein (Feedipedia 2012). The resulting flour could be used textured or as is for incorporation into many preparations, provided a white color is not desired.

<figure><img src="image_13.png" /></figure>
This product is offered in the “organic agriculture” sector in Germany (AOT 2023). A more concentrated version, delipidated by supercritical CO₂, is also available with a less pronounced color.

### 3.3 Impact of genotypes (G), environments (E), cultural practices (P), and interactions (G×E×P)

The synthesis of SSP is primarily regulated at the transcriptional level (Xi and Zheng 2011), but abiotic stresses, whether climate (elevated atmospheric CO₂ concentration, drought, high temperatures...) or plant nutrition (nitrogen or sulfur deficiencies), strongly influence seed protein accumulation and composition in crop species (Bonnot et al. 2017; Bourgeois et al. 2009; Chandler et al. 1984; D'Hooghe et al. 2014; Dupont and Altenbach 2003; Tabe et al. 2002). For example, in soybean, water stress affects the amount of protein per seed but does not decrease the protein concentration because the oil is also significantly reduced (Rotundo and Westgate 2009). Water deficit leading to stomatal closure reduces photosynthesis and, thus, starch accumulation, especially in wheat. It accelerates leaf senescence and the remobilization of nitrogen from the leaves to the grains. Depending on the intensity, duration, and other characteristics of the stresses experienced by the plant, the protein concentration of the grain can sometimes be increased (Sehgal et al. 2018). Significant genotype × environment interactions have been observed for protein content in soybean (Whaley and Eskandari 2019), lentil (Subedi et al. 2021), pea (Klein et al. 2020), and wheat (Bnejdi and Gazzah 2010) species.

Understanding the genetic basis of these interactions would provide information about the genes that contribute to phenotypic plasticity, i.e., the ability of a genotype to change its phenotype according to environmental parameters. A study conducted in *Medicago truncatula* suggests that globulin plasticity in response to the environment relies on the plant's ability to recycle methionine (Cartelier et al. 2021). Compared to other abiotic stresses, sulfur deficiency has a strong negative impact on seed protein composition since it massively decreases the proportion of S-AA-rich proteins (Bonnot et al. 2017; D'Hooghe et al. 2014; Henriet et al. 2019). The change in SSP composition in wheat depends on the balance between nitrogen and sulfur fertilization (Bonnot et al. 2020; Yu et al. 2021). In this species, the decrease in S-AA modifies the flour health quality (for instance, S-rich gliadins containing the immunodominant epitopes involved in coeliac disease) and its end-use quality (for example, bread-making potentialities) (Zhao et al. 1999). Candidate genes for the response of developing seeds to the sulfur deficiency have been identified using “omics” approaches in wheat and pea (Bancel et al. 2019; Bonnot et al. 2020; Henriet et al. 2019, 2021). In addition, Cartelier et al. (2021) show that one of these candidate genes

encodes a homocysteine S-methyltransferase3 (HMT-3), which regenerates methionine from S-methylmethionine and contributes to the accumulation of 11S globulins even under sulfur-deficient conditions. Similar results have been reported in wheat (Yu et al. 2021). These authors also showed that adequate sulfur supply increases grain protein yields through improved nitrogen use efficiency. In addition, the role of an HMT (HMT-1), whose expression is regulated by a transcription factor belonging to the MYB family and controlled by sulfur availability, has been reported (Yu et al. 2021). Adequate sulfur intake leads to overexpression of genes encoding myb factor and HMT-1, which may promote threonine and glycine biosynthesis in favor of glutenin synthesis (Yu et al. 2021).

The literature is scarce regarding the genetic and molecular determinants of plasticity in seed protein content or composition. A better understanding of these determinants would help reduce the impact of environmental variations: a research effort in this direction would be welcome. Furthermore, in the context of the agroecological transition, it is essential to assess the impact of low-input cropping practices (and the shift from synthetic to organic nitrogen sources) on seed protein content, composition, and quality and to determine how intercropping and diversification in crop rotations contribute to this quality.

### 3.4 Crop associations: in spite of many advantages, disruptive cropping practices slow their adoption

Crop associations can decrease significantly the needs for synthetic inputs while producing almost as much as their best component (Li et al. 2023). From 2005 to 2010, studies based on multi-partner trials highlighted the interest of wheat (bread or durum)/legume (protein pea, chickpea) associations on the quantity and quality of the harvest (Bedoussac et al. 2015). Indeed, legumes acquire nitrogen essentially thanks to their nodules that fix atmospheric nitrogen due to the presence of specific bacteria (nitrogen-fixing rhizobia). Wheat grown in association with a legume species has a reduced number of ears and grains/m² compared to a monoculture, but its grain yield is only slightly reduced (the plot yield is even higher if we consider the double crop wheat + pea) (Bedoussac et al. 2015; Li et al. 2023). An increase in grain protein levels without the additional use of mineral nitrogen is also observed. Indeed, the nitrogen functioning of the plot is improved because of the nitrogen competition at the root level: wheat is a strong competitor and acquires more resources, the legume compensating by an increased activity of symbiotic nitrogen fixation (Bedoussac et al. 2015; Jensen et al. 2020; Hauggaard-Nielsen et al. 2008; Li et al. 2023). The effect of cereal-legume intercropping on cereal protein content is well known, but
to date, no breeding program has addressed the develop-
ment of varieties with high mixing ability, despite the known
changes in performance and ranking of varieties grown
in single-cropping versus intercropping (Annicchiarico
et al. 2019, Haug et al. 2023, Demie et al. 2022, Moutier
et al. 2022). Crop associations also improve production resil-
ience in the context of climate hazards frequency increase,
resulting from climate change. This ability is derived from
the ecological principles of complementarity, cooperation,
competition, and compensation between crops, the so-called
“4C approach” (Bedoussac et al. 2015; Justes et al. 2021).
Note that to a lesser extent, variety mixtures can also confer
advantages regarding protein composition. For example, in
bread wheat, work carried out within the framework of the
Wheatamix project (Barot et al. 2017; Borg et al. 2018) has
shown that variety mixtures allow a modest gain in yield,
can contribute to improving bread-making quality, and erase
the defects of one component of the mixture. In practice,
crop associations do present advantages and should be inte-
grated into rotations.

The balance between associated crops is challenging to manage because the outcome of their competition is highly dependent on environmental factors (soil fertility, climate, diseases...), and the proportions of species harvested can vary significantly between 2 years or two sites (Gaba et al. 2015). Recent meta-analyses showed yield gains of about 20% in these interspecific associations compared to monocultures (Bedoussac et al. 2015; Beillouin et al. 2021; Cernay et al. 2018). Since cereals are generally more competitive with weeds than legumes, they typically provide better control than legume monocultures. Finally, disease and weed pressure are generally reduced. However, the concordance of maturity of wheat and legume grains for harvesting is challenging to manage, and sorting grains after harvest is a constraint. Thus, despite the interest in these practices, they are not widely implemented.

However, use of crop associations is increasing, as
observed in sustainable agriculture, both from farmers'
initiatives and thanks to projects that have promoted their
development (e.g., H2020 SolACE project (2025), H2020
ReMIX project (2023), DIVERSify project (2023) and
MoBiDiv project (2023)). This adoption of associated
cereal/legume crops, particularly with lentils (see https://
www.fermiersdeprovence.fr/, https://www.duransia.coop/),
is due to several reasons (Jezequel 2021):

* Market demand: there is a strong demand for lentils from France, produced in a “short chain,” preferably in organic culture, with a remunerative price disconnected from the global price.

* Removal of a technological barrier: most grain storage agencies are now equipped with high-capacity optical sorters for grain/legume separation.

* Creation of diversification chains, going all the way to the consumer: despite the administrative and commercial logistical constraints, cooperatives and traders, even of small size, are evolving towards the downstream part of the chains to pass on the added value to the producers.

* A complete change of vision of the producers regard-
  ing the benefit of the association technique: lentil has
  become profitable, whereas the research world initially
  imagined cereals as the main crop. The beneficial effects
  of the association for lentils are observed: the cereal
  acts as a “stake,” which facilitates harvesting, occupies
  space, and reduces weed pressure. The profitability of the
  production depends on the success of the lentil, and the
  harvested wheat constitutes an additional income.

4 From proteins to foods: designing food products adapted for diverse consumer needs and demands

## 4.1 Optimization of protein digestibility for human consumption

Cereal proteins have good digestibility: around 90% of wheat flour in humans (Bos et al. 2005). Digestibility is generally lower for whole grain (Han et al. 2019), as bran, fiber, and phytates interfere with proteins and decrease protein access. Similarly, pulses, such as peas and soybeans, have highly digestible proteins (90% at least) in the form of isolates or concentrates (Gaudichon et al. 2002; Guillin et al. 2022; Hertzler et al. 2020; Mariotti et al. 2001). When consumed as seeds, their digestibility decreases and reaches 60 to 85% depending on the species and the preparation. For example, the standardized ileal digestibility in pigs of six faba bean cultivars is between 70 and 81% (Jezierny et al. 2011). As for cereals, fiber and anti-nutritional factors are responsible for this low digestibility. In particular, β-glucans form aggregates with lentil and pea proteins, strongly decreasing their digestibility in vitro (Boachie et al. 2021). Some practices, notably dehulling and extrusion, can modulate the digestibility of dry vegetable proteins (Nosworthy et al. 2018) and increase the digestibility of mung bean amino acids by 6% (Devi et al. 2018) and chickpea amino acids by 15% (Devi et al. 2020). Soaking and cooking methods are also levers to modulate the content of phytates, tannins, and trypsin inhibitors and, consequently, the amino acids' digestibility (Badjona et al. 2023; Gilani et al. 2012).

There is still a lack of knowledge about the digestibility
of proteins in specific populations. In stunted children (Devi
et al. 2018), the digestibility of amino acids seems to be
higher than in adults (Devi et al. 2020), but this remains to
be proven by a dedicated study. Protein bioavailability in
older adults is a significant issue in the context of the shift

<figure><img src="image_14.png" /></figure>
to a high proportion of proteins from plants in the diet of the aging population. While protein intake is often below requirements in this population, digestibility and, more broadly, protein quality are crucial issues. While the effect of age on intestinal motility and pancreatic secretions is well documented (Rémond et al. 2015), the consequences on digestive efficiency remain largely unexplored in humans and animal models. Only one study in young and 20-month-old rats (Gilani and Sepehr 2003) shows that protein digestibility decreases with age, especially for less digestible foods, including raw or cooked soybeans or beans. Still, age has a low impact on the digestibility of already highly digestible proteins, such as those from milk.

## 4.2 Attractiveness of plant proteins for market development

An increase in the proportion of plant proteins in our diets implies an increase in the consumption of pulses, cereals, and vegetable substitutes for animal products. While cereals are already an essential part of our diet, the supply of pulses or foods enriched with plant proteins must be increased. Similarly, it is possible to develop the uses of soybeans in traditional Asian forms by remedying their drawbacks. Based on our analysis of the literature, five main possibilities can be considered depending on the type of food and processing in terms of tradition or innovation (Table 2):

(1) Modify traditional processing to improve pulses consumed traditionally.

(2) Develop innovative foods through traditional processes (e.g., fermentation).

(3) Adopt “exotic” products, possibly adapted to local tastes and culinary practices.

(4) Adapt processes for mixing pulses and cereals in traditional foods.

(5) Use recent technologies (e.g., extrusion) to develop new foods.

Whichever pathway is followed, the process will inevitably modify the complex structure of the grain, which is composed of the pericarp (a series of envelopes), a reserve tissue

(endosperm in cereals or cotyledons in legumes, where starch, proteins, or lipids accumulate), and the germ of the future seedling. The envelopes of pulses are covered with wax and are, therefore, less permeable than those of cereals. The endosperm of cereals consists of cells surrounded by hemicellular walls. At the level of the cotyledon, the reserve compounds are stored in cells whose arrangement allows water circulation. For cereals and pulses, the water accessibility by capillarity is favored by compartments like the hilum or germ.

### 4.2.1 Modifying processing to improve pulses consumed traditionally

The preparation and processing of pulses can be an obstacle to their consumption. One of the main obstacles to the traditional consumption of pulses by households is the time required for their preparation, which often requires a soaking step to hydrate the seed, eliminating some of the anti-nutritional factors. Some food companies perform a hydrothermal pretreatment of the seeds by precooking at temperatures high enough to promote hydration and initiate swelling of the starch granules to reduce the preparation time in households. The temperature of this pretreatment depends on the water content of the raw seed and can be predicted from a state diagram (Figure 4). Since heat diffusivity is higher than water diffusivity, around $10^{-7}$ versus $10^{-10}$ m$^2$/s respectively, grain hydration is the limiting mechanism for cooking and germination (Miano and Augusto 2018). Hydration kinetics can be controlled and modeled, but the role of different entities (cell walls and reserve tissues) in water diffusion is still unknown. Hence, the control of pretreatment of pulses requires further studies.

### 4.2.2 Developing new foods through traditional processes (e.g., fermentation)

Pulses are used in a broad range of food products, with various processes, to replace casein and whey protein in fermented and unfermented dairy products, to be used as substitutes for animal protein, or to enrich protein levels in baked foods, cereals and snacks (Ben-Harb et al. 2018; Monnet et al. 2022; Petitot et al. 2010; Pinel et al. 2023). Indeed,

<table><caption>**Table 2** Main tracks for the development of pulses-based foods. The numbers refer to the paragraphs of Section 4.2</caption><thead><tr><th>Process</th><th>Food<br>Traditional</th><th>New</th></tr></thead><tbody><tr><td>Traditional</td><td>1. Precooked</td><td>2. Analogs of traditional dairy foods using plant protein isolates</td></tr><tr><td>Traditional exotic</td><td>3. Tofu, hummus, tempeh and their “pea” or “lentil” variations</td><td></td></tr><tr><td>Adapted or new</td><td>4. Pulses/cereals mix</td><td>5. Meat analogs</td></tr></tbody></table>

<figure><img src="image_15.png" /></figure>
<figure><img src="image_16.png" /><figcaption>**Fig. 4** A primer to identify processing conditions: state diagram representing variations in melting and denaturation temperatures of wheat (orange) and pea (green) starch and protein with water content. Tm indicates melting temperature and the curve with orange dots represents the variations of the temperature of melting onset. These variations are supplemented by thermal routes (dotted lines) of different processes reported in Table 2: precooking (1), fermentation (2), e.g., cooking of cereal/legume mixtures (4), and meat analogs by wet cooking-extrusion (5). Source: Della Valle et al. (2019) and Mitchell et al. (1994)</figcaption></figure>

pulse proteins are good candidates for designing healthy and sustainable foods due to their interesting and various functional properties.

For this purpose, legume seeds can be processed into flour, concentrates, and isolates. These ingredients are obtained by air-drying, dehulling, grinding of seeds, and protein extraction, which may undergo further modification by appropriate processes and conditions. However, the use of these ingredients can be hindered by sensory defects: an unpleasant smell and taste, often related to a green, herbaceous, or bean-like aroma (notably attributed to aldehydes, alcohols and ketones derived from the action of lipoxygenase), but also bitter and astringent flavors potentially contributed by sapid compounds such as saponins and phenolic compounds (isoflavones, flavonols, tannins, hydroxycinnamic acids, among others) (Sharan et al. 2021).

The fermentation process of plant-based foods, using various and multiple microorganisms used classically for fermented dairy products, constitutes a very interesting lever to develop innovative food products. For example, lactic acid fermentation can improve the taste and odor of pea-based products by reducing or masking the unpleasant “green note” off-flavor. Thus, pulse proteins could expand the sources of nutrients subject to proteolytic microorganisms. These proteins may or may not be mixed with milk

and fermented by various microbial consortia to modify flavor compounds, including those responsible for the “green note,” with implications for consumer perception (Ben-Harb et al. 2020; Saint-Eve et al. 2021). Recently, this work has led to the development of a cheese analog based on pulse proteins called “camenvert” (Figure 1). Nevertheless, the interactions between microbial consortia and the adaptive metabolism of microorganisms need to be studied, combining sensory, biochemical, physiological, and meta-omics analyses to control their growth in plant protein gels better.

### 4.2.3 Adopting “exotic” products

Traditional processes developed in Southeast Asia, such as fermentation (e.g., tempeh from soybeans (cooked and dehulled soybeans, fermented by a mold of the genus Rhizopus)), are an exciting lever to overcome these sensory defects and develop food products with diversified properties. There is a wide variety of soy foods, from the simple gel (tofu) to whole beans (natto) to fermented gels (e.g., furu, misozuke, chao, and others), with typical tastes and aromas. However, these products are unfamiliar for their texture or taste, slowing their adoption. Therefore, recipes adapted to French or European culinary practices and palates may need to be invented or popularized. In the case of

<figure><img src="image_17.png" /></figure>
soybean derivatives, the preparation method influences the isoflavone content (Fernandez-Lopez et al. 2016). Ultrafiltration reduces the isoflavones/protein ratio from 6.5 to 1.2. Fermentation of tonyu by microorganisms equipped with α-galactosidases minimizes the presence of oligosaccharides (Hati et al. 2014).

### 4.2.4 Adapting processes for mixing pulses and cereals in traditional foods

Agricultural research has demonstrated the potential benefits of cereal/legume combinations (see Section 3.4). There is a genuine interest in promoting the use of mixed flour that would favor the elaboration of foods enriched with vegetable proteins. However, the rheological properties and processing ability of cereal flours are modified by adding pulse flours. This result has been illustrated in the particular case of the manufacture of initially cereal-based foods (Bresciani and Marti 2019; Monnet et al. 2019), such as snacks (Kristiawan et al. 2021), pasta (Berrazaga et al. 2020a, 2020b), cookies, cakes (Monnet et al. 2022), and bread (Atudorei and Codina 2020), where structural modifications result in textual changes. Tools and concepts such as biopolymer state diagrams will help predict gelatinization and denaturation temperatures of starch and protein, respectively, as a function of water content leading to the development of processes for producing cereal products enriched with pulses (Figure 4). In this sense, it will be possible to make the necessary processing adjustments to compensate for the dilution of the gluten network in the case of bread or pasta, improve the aeration of sponge doughs, or modulate the viscosity and melt expansion of snacks made by dry extrusion (Monnet et al. 2019). While the adverse sensory effects inhibit the consumption of these foods, the appropriate flavor formulations can compensate for them.

### 4.2.5 Using new technologies (e.g., extrusion) to develop new foods

The demand for plant-based “meat analogs” as an alternative food illustrates the value of technology in developing novel foods. Meat analogs, relatively inexpensive compared to traditional meat, are strictly plant-derived products, often made with plant protein isolates (Boehm et al. 2023; van der Sman and van der Goot 2023). They are designed to have a texture similar to meat using a high-water content cook-extrusion process (Cheftel et al. 1992; Guyony et al. 2022). Texturizing vegetable proteins using this process involves an initial step at temperatures above 120 °C (Figure 4), where the proteins are denatured by the shear imposed by the rotation of the extruder screws. Subsequently, the formation of bonds between the denatured protein chains, such as disulfide or

hydrogen bonds, starts near the extruder exit. As the protein “melt” flows through the extruder die, the axial flow creates a fibrous structure by aligning the cross-linked peptide chains (Ryu 2020). Textured products obtained by wet extrusion are not dried; they are refrigerated and can be consumed directly. Thus, modifying the extrusion parameters can influence the structure and texture. However, work is still needed to improve sensory properties and to verify that denaturation (i.e., unfolding and refolding of proteins) does not decrease the nutritional quality of the seeds, although extrusion would have a positive effect on the digestibility of chickpeas (Devi et al. 2020).

A better understanding of the structural modifications of foodstuffs at the different levels of matter organization during the process would enable manufacturers to better satisfy consumer expectations. Moreover, producing ingredients based on pulses makes it possible to consider substitutes for food additives (Burger & Zhang 2019), in line with the “Clean Label” claims, for less processed and healthier products. However, regardless of the processing route considered, foods (including ingredients) rely on a stable supply and selection of plant raw materials.

## 4.3 Algae and microalgae, a niche with added value

The latest trends for plant protein production include macroalgae (Juul et al. 2024, Siladji et al. 2024), while in recent years, microalgae have been developed as new sources of proteins of interest. However, the term “microalgae” includes eukaryotic monocellular algae (i.e., plants, photosynthetic and with plant cell walls) but also cellular prokaryotic microorganisms (no nucleus or organelles), which are very distant from a phylogenetic point of view. Because these share key characteristics and are confounded in common language, both will be described here.

Compounds from algae contain many nutrients and constituents of interest (proteins, fatty acids, carotenoids, or vitamins). Microalgae have attracted attention due to their high protein content, reaching 70% of the dry biomass (Matos 2019). The FAO has recommended spirulina production for over 20 years (Habib et al. 2008) to help meet the need for food supplements and animal feed in underdeveloped countries. As another example, oligo-polysaccharides from microalgae are considered candidates for prebiotic properties (Patel et al. 2021).

Microalgae production has been expanding over the last decade with a 150% increase in the number of companies involved in Europe (Araújo et al. 2021) for a wide range of applications adding value to food, nutraceuticals, cosmetics, and pharmaceuticals. The human food markets use most of the microalgae biomass produced in the world, estimated at 20,000 tons per year. This production is increasing by about 10% every year (SABANA 2018). Microalgae accepted for human food in Europe are listed in Table 3 (CEVA 2019).
<table><caption>**Table 3** Microalgae approved for food use in the European Union</caption><thead><tr><td>Species, biological entity</td><td>Food applications</td></tr></thead><tbody><tr><td>Aphanizomenon flos aquae Ralph*</td><td>No restrictions</td></tr><tr><td>Arthrospira sp. (spirulina)*</td><td>No restrictions</td></tr><tr><td>Arthrospira platensis Gomont (spirulina)*</td><td>No restrictions</td></tr><tr><td>Chlorella sp.</td><td>No restrictions</td></tr><tr><td>Chlorella luteoviridis Chodat</td><td>No restrictions</td></tr><tr><td>Chlorella pyrenoidosa Chick</td><td>No restrictions</td></tr><tr><td>Chlorella vulgaris Beijerinck</td><td>No restrictions</td></tr><tr><td>Haematococcus pluvialis Flotow</td><td>Condiments</td></tr><tr><td>Odontella aurita Lyngbye**</td><td>Condiments in finished products</td></tr><tr><td>Tetraselmis chuui Butcher</td><td>Sauces, condiments</td></tr><tr><td>Schizochytrium sp. extracts</td><td>Oil</td></tr><tr><td>Ulkenia sp. extracts</td><td>Oil</td></tr></tbody></table>

Other than those listed in Table 3, microalgae are authorized by the French authorities only for nutraceuticals (food supplements). Their high content of proteins and other nutrients, including B vitamins, iron, and zinc, makes them attractive species for biological applications and/or for improving the nutritional quality of food. *Chlorella* sp. and *Arthrospira* sp. proteins can potentially reduce the risk of anemia (Mohamed et al. 2013). Spirulina is believed to have many health benefits: its proteins are thought to improve the immune and lymphatic systems and provide additional protection against ulcers. As for those from *Arthrospira maxima* (Setchell & Gardner), they could reduce the risk of ulcers, constipation, anemia, hypertension, diabetes, and infant malnutrition (Camacho et al. 2019). Phycocyanin, a specific phycobiliprotein extracted from microalgae, has already been used in food supplements and the food industry as a blue dye and for its biological properties of interest: it has antioxidant, anti-inflammatory (Romay et al. 2003), antiallergic (Remirez et al. 2002), and immune activities (Capelli and Cysewski 2010; Khan et al. 2005).

Proteins isolated from microalgae have an amino acid profile that gives them good nutritional quality (Černá 2011). Two species are particularly rich in arginine, *Nannochloropsis salina* Hibberd and *Dunaliella salina* Teodoresco, as this amino acid contributes to the limitation of cardiovascular diseases. Marine microalgae are known to have a more balanced amino acid profile than freshwater algae (Nguyen et al. 2020). Microalgal proteins show in vitro digestibility of 50–80% depending on the source, model, and method used: the digestibility of non-extracted proteins from *Chlorella vulgaris* Beijerinck, *Phaeodactylum tricornutum* Bohlin, *Arthrospira platensis* Gomont, and *Nannochloropsis oceanica* Suda & Miyashita reaches 79%, 77%, 74%, and 54%, respectively (Wild et al. 2018).

Thus, although they have interesting amino acid profiles, their digestibility and nutritional quality remain lower than animal or some plant protein sources for which digestibility is above 90%. It can be improved by extraction, which makes them more easily accessible to digestive enzymes (Barros de Medeiros et al. 2022). In vivo studies indicate protein digestibility values of 82% for whole spirulina in rats (Tessier et al. 2021) and 86% in humans (Devi et al. 2020). Some microalgae may have promising functional properties (emulsifying, stabilizing, foaming) for human food applications (McCarthy et al. 2013).

However, microalgae production still faces significant challenges before it can be fully integrated into food systems and not only as supplements.

1. Price is the first issue. For example, 1 kg of dry spirulina biomass grown in open ponds, with a protein content of 70%, cost between 3 and 10 € in the early 2010 (Delrue et al. 2017). The price varies according to the type of microalgae (SABANA 2018) and the type of reactor used. The prices range between 10 and 100 $/kg for production in open ponds and 30 $/kg for production in photobioreactors. In addition, the cost of extraction of some molecules must be considered.

2. Secondly, not all production processes can be scaled up to an industrial scale in terms of yield, cost, or tooling (Matos 2019).

3. Thirdly, organoleptic characteristics remain an issue (Sidari and Tofalo 2019). Processes are being developed to produce microalgae ingredients with less odor, color, and taste, such as those from *Chlorella vulgaris* (marketed by Allmicroalgae).

4. Another limitation is the potential risk of adverse reactions due to peptides or toxic compounds from microalgae (Hadi & Brightwell 2021).

5. Lastly, microalgae hydrolysates may need to be chemically treated to ensure their safety and compliance with regulatory standards (Apone et al. 2019).

In the European Union, new strains of microalgae or their extracts intended for human consumption fall under the regulatory framework of the Novel Food directive. Some microalgae (*Nannochloropsis gaditana* Lubián, *Euglena gracilis* Klebs) are currently being examined for validation through this procedure.

Seaweeds (macroalgae) have the advantages of not competing for land or water use and of being already industrially produced. Although some of the more common algae used in food such as nori (*Porphyra purpurea* Roth used for sushi) or wakame (*Undarai pinnatifida* Harv.) are rich in proteins (Taboada et al. 2013), they are also rich in polysaccharides which hinder access and digestibility of their proteins (Juul et al. 2024). Consumption of seaweeds as such appears likely

<figure><img src="image_18.png" /></figure>

<footnote>*Cyanobacteria. **: Diatom</footnote>
to stay limited in Europe due to their strong sensory impact, i.e., use as seasoning or for functional properties (Siladji et al. 2024) rather than bulk food, and extraction of proteins from algae is still in its infancy.

# 5 General discussion and identification of research priorities

From our knowledge of the existing literature and exchanges, notably during a 2-day workshop entitled “Protéines végétales pour la santé des consommateurs et des systèmes de production” (Plant proteins for the health of consumers and production systems), several research needs and their interdependencies could be identified at all value chain stages (Figure 2). This paragraph presents the results of this critical analysis by the authors of the scientific literature and their discussion and confrontation.

## 5.1 An urgent need of connecting various disciplines and points of view

There are numerous levers for improving the quantitative and qualitative supply of plant proteins, from which various pathways are identified, for instance for legumes development (Cusworth et al. 2021b). Their implementation must be built on a transdisciplinary basis allowing them to couple the issues of upstream production, genetics, agronomy and downstream transformation, process, biochemistry, microbiology, sensory, and nutrition, among others. Understanding the barriers to their adoption requires mobilizing human, social, and economic sciences to identify the determinants of behavior from grower-producers to consumers, not forgetting processors, and initiating future innovations towards their success. Thus, multi-dimensional and multiactor approaches are to be favored to integrate the possible constraints and levers, facilitating the development of a new offer that contributes to the transition of our food systems.

In this context, cultural practices, species, varieties, and strains must be studied and renewed to reach the double objective of being compatible with the production chains from farm to plate and reducing input use. This paper shows the need for the integration of a broad range of crops' characteristics to meet the various users' expectations. It starts with the characteristics of the varieties to be bred to fit with the growers' cropping systems and the expected use of the harvests. To reach and integrate these expectations, the process of ideotype variety building is major (Donald 1968). The ideotyping process consists of defining the ideal characteristics of a variety best adapted to defined/expected growing conditions and harvest use (Debaeke et al. 2014). The visions of breeders, producers, and processors can be combined through an approach such as the varietal ideotyping

process in order to integrate the anticipated difficulties. The food sciences are involved throughout the supply chain. They must determine the nutritional and organoleptic quality of the desired products and ensure that this quality is maintained or improved during the transformation process. An essential need is to better link upstream and downstream, from breeding to processing, to optimize the whole process. For example, if anti-nutritional factors are eliminated by sustainable processing, it will not be necessary to counter-select against them during variety creation.

There is a proven difficulty in turning an innovative idea into a prototype or product in the marketplace. This challenge is reflected in the critical transition from the upstream stages of the TRL (Technology Readiness Level) scale to the downstream stages of development, commonly referred to as the “valley of death” of the TRL scale. This complicated passage consists of proof-of-concept implementation and project maturation. Different tools are available today to help overcome these difficulties.

Furthermore, we are observing a critical period that allows us to combine many cognitive and methodological achievements: translational genomics, high-throughput sequencing and phenotyping, association genetics, and predictive approaches, among others. Wise tool combinations offer opportunities to accelerate genetic progress on minor species substantially. The success of such approaches will also depend on varietal and crop ideotyping work that considers the expectations of the downstream sectors to ensure good valorization for harvested products while considering the challenges that production must meet to support the transitions underway. The upstream-downstream dialog must be intensified, particularly within the CTPS.

## 5.2 Interdisciplinary research needs

The following interdisciplinary research priorities identified in our working group to help overcome the resistance to change and increase protein-rich plant production and use could be formulated and related to Figure 2. All of these research needs, though addressing more specifically one of the different sections in Figure 2, require an interdisciplinary and concerted effort. Block 2 and 4 for example are mirror images for proteins, concerning for block 2 their production in the plant and for block 4 their resulting nutritional qualities.

* Interdependancies:

- Explore and evaluate organizational arrangements and incentive levers that promote shared dialog between upstream and downstream sectors to support joint innovation trajectories for the resilience of agri-food systems.
- Study and understand consumers' wishes, expectations, and behaviors and the possible adaptation of new products to the habits of food use.

- Promote transdisciplinary and concerted studies between geneticists, agronomists, and biochemists to characterize the links between cultivation practices, plant breeding, transformation processes, and optimized protein compositions for human and ecosystem health.

* Block 1 “diversity”

- Evaluate the effect of agroecological cropping itineraries, particularly those mobilizing diversification at different time and space scales, on seed protein content, composition, quality, and stability over environments.

- Identify and characterize the current technological challenges, in particular, those associated with agricultural equipment, that make it possible to enhance the value of heterogeneous harvests, with variable qualities, from crops grown in mixtures and polyculture.

- Develop new protein sources for food, such as underutilized and locally produced legumes and pulses, oilseeds, pseudocereals, and microalgae, to diversify the offer in terms of quality.

* Block 2 “composition and yield”

- Define new ideotypes to identify and characterize the traits of the plants to be selected to produce protein-rich foods for human and animal health and the environment.

- Explore and identify ways to improve the amino acid composition of grains for human consumption, using genetic and agronomic levers, without increasing nitrogen or sulfur fertilization.

- Develop breeding programs favoring diversification, through the development of varieties adapted to new cropping systems, as crop mixtures, and new species of legumes and pulses.

* Block 3 “processes”

- Conceive processes for the sober production of new products accepted by the consumers, endowing value to heterogeneous harvests with variable qualities.

- Understand foods' structural, sensory, nutritional, and functional modifications during domestic and industrial preparation processes, including using microorganisms.

* Block 4 “nutritional qualities”

- Document the specificities and complementarities of the nutritional qualities of crop species, including minor species, to guide crop diversification towards more resilient food systems.

- Explore and identify the levers for improving the digestibility and/or organoleptic properties of plant foods (cereals, pulses, and others) and the bioavailability of their nutrients.

Since the inception of the interactions between researchers and identification of the macrotopics at different steps in Figure 2, the French government launched two research calls on alternative proteins. In this context, the projects listed in Suppl Table, which benefited from the interactions initiated by Carnot Qualiment, Carnot Plant2Pro, and PlantAlliance, were funded. Many of these projects are interdisciplinary, adopting a farm-to-fork approach and focusing on the organizational levers as well as technical levels.

## 6 Conclusion

The aim of this review was to bring together disciplines from economics to nutrition to identify the research needed to enable the necessary systemic innovation for rebalancing of protein intake between animal and plant proteins in healthy and sustainable diets. It was also to help actors of transfer, such as Carnot Qualiment, Carnot Plant2Pro, and PlantAlliance, identify priority topics for support and transfer to socioeconomic actors.

Given the challenges for global health, particularly the health of the environment and human health, significant research needs to be conducted to make vegetalization of protein intake (and therefore of diets) operational. The crucial issue is that of interdependencies in the food system between production, processing, and consumer acceptance, which will require transdisciplinary and concerted studies. Two further points which our review highlights are (i) identifying the balance between plant and animal production and (ii) integrating other components beneficial to health, such as polyphenols or polyunsaturated fatty acids, in the discourse about vegetalization of diets. In this context, it seems urgent to devote a significant part of public research and support for innovation to this rebalancing, as its funding lags well below that devoted to synthetic proteins or in vitro meats, whose benefits in environmental and social terms are disputed.

The research work identified in this review must be undertaken in the short term and on time steps compatible with the marketing in the medium term of new varieties and new foods, whereas environmental impact studies must be planned in the

<figure><img src="image_19.png" /></figure>
long term. The importance of the issue and the scope of the work for the common good must mobilize public and private actors with strengthened interactions to promote innovation.

**Supplementary Information** The online version contains supplementary material available at https://doi.org/10.1007/s13593-025-01034-1.

**Acknowledgements** Co-first authors Karine Gallardo-Guerrero and Anne Saint-Eve contributed equally to the manuscript. The authors thank Denis Chéreau (Improve), Frédéric Muel (Terres Inovia), Isabelle Solanas (GreenTech), Maxime Szambien (PlantAlliance), Vanessa Verdoud (INRAE), and Stéphane Walrand (INRAE) who contributed to the whole reflection. We thank Christine Le Signor (UMR Agroécologie) for her review of the document and her constructive remarks concerning the genetic determinism of the protein value of pea seeds studied in the framework of the French National Research Agency (ANR)-PEAV-ALUE project (ANR-19-CE21-0008). We also thank ReadyToPub- Author Services Provider for an efficient and constructive service. The projects cited in Suppl Table are financed by ANR-France 2030.

**Authors'contributions** All authors contributed to the final manuscript's writing, revision, and editing and approved the final version for submission; RC, MDF, and CMGCR designed the review and carried out the final editing of the manuscript.

**Funding** This study was funded by PlantAlliance (publishing fees) and INRAE Dept TRANSFORM (translation and English revision fees).

**Data availability** Data sharing is not applicable to this article as no datasets were generated or analyzed during the current study.

**Code availability** Not applicable.

## Declarations

**Ethical approval** Not applicable.

**Consent to participate** Not applicable.

**Consent for publication** Not applicable.

**Conflict of interest** The authors declare no competing interests.

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