The Journal of Nutrition

Nutritional Epidemiology

Approximately Half of Total Protein Intake by

Adults Must be Animal-Based to Meet
Nonprotein, Nutrient-Based
Recommendations, With Variations Due to
Age and Sex
Florent Vieux,1 Didier Rémond,2 Jean-Louis Peyraud,3 and Nicole Darmon4

1
MS-Nutrition, Marseille, France; 2 Université Clermont Auvergne, INRAE UNH, Clermont-Ferrand, France; 3 UMR INRAE-ACO
Pegase, St Gilles, France; and 4 MoISA, University of Montpellier, CIRAD, CIHEAM-IAMM, INRAE, Institut Agro, IRD, Montpellier,
France

ABSTRACT
Background: Shifting towards a more plant-based diet, as promoted in Western countries, will reduce the animal
protein contribution to total proteins. Such a reduction may not only impair protein adequacy, but also the adequacy in
other nutrients.
Objectives: We determined, for different adult subpopulations, the minimum total protein levels and the minimum
animal protein contributions to total proteins that are compatible with the fulfillment of all nonprotein nutrient-based
recommendations.
Methods: Mean nutritional contents and mean diet costs were estimated using a French, cross-sectional,
representative survey for 5 French subpopulations: 1) women < 50 y; 2) women 50–64 y; 3) women ≥ 65 y; 4) men < 65
y; and 5) men ≥ 65 y. For each subpopulation, linear programming optimization was used to assess the minimum protein
level (model set #1) and the minimum animal protein contribution to total proteins (model set #2) that are compatible
with the fulfillment of all nutrient-based recommendations (except proteins, for which levels were analyzed as outputs).
Total diet costs were not allowed to increase. Eating habits were considered in model set #2 only.
Results: The minimum amount of protein that was theoretically compatible with the fulfillment of nutrient-based
recommendations (model set #1) was below the minimum recommended protein intake for all subpopulations except
women < 50 y. In model set #2, for women and men ≥ 65 y, decreasing animal protein contributions to total proteins
below 55% and 60%, respectively, led to protein levels below recommended levels. For the other subpopulations
(women < 50 y, women 50–64 y, and men < 65 y), the lowest animal protein contributions to total proteins compatible
with a nutritionally adequate diet (including protein adequacy) were 55%, 50%, and 45%, respectively.
Conclusions: This study provides factual information about the animal protein contributions to total proteins compatible
with meeting all nutrient-based recommendations at no additional cost, and shows that they vary between 45% and
60% depending on the group of adults considered. J Nutr 2022;152:2514–2525.

Keywords: animal-to-plant protein ratio, nutritional adequacy, optimization, protein quality, affordability, diet cost,
France

Introduction (1). Higher consumption of plant-based foods has consistently

been associated with positive health outcomes (2), whereas
Guiding principles for sustainable and healthy diets include excess consumption of red meat and processed meat is often
eating an abundance and a variety of minimally processed, discouraged by public health nutrition guidelines (3).
plant-based foods (fruits, vegetables, nuts, unrefined cereals, Because meat has a higher protein content per kilocalorie
legumes, etc.) and moderate amounts of animal-based foods than plant-based foods, shifting towards more plant-based diets


C The Author(s) 2022. Published by Oxford University Press on behalf of the American Society for Nutrition. This is an Open Access article distributed under the
terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and
reproduction in any medium, provided the original work is properly cited.
Manuscript received April 6, 2022. Initial review completed June 13, 2022. Revision accepted July 1, 2022.
2514 First published online July 11, 2022; doi: https://doi.org/10.1093/jn/nxac150.
and less meat will inevitably reduce both total dietary proteins B6), minerals (magnesium, zinc), and water differ for men and
and the animal protein contributions to total proteins (4). women (Table 1). Then, for each sex, an age threshold of 65
Animal proteins have a more balanced indispensable amino acid y was used in order to take into account the differences in
profile and greater digestibility than plant proteins, although recommended levels of protein intake: 1.0 g of proteins/kg
plant proteins are commonly depicted as being of lower of body weight for adults aged 65 years old and older and
quality than animal proteins (5). Yet, in Western countries, 0.83 g of proteins/kg of body weight for younger adults (7).
where the quantities of protein consumed generally exceed The subpopulation of women was further divided according
minimum requirements, quality differences between proteins to iron needs, considering 50 y as the menopausal year (13).
from different food sources have negligible impacts on protein This led to 5 subpopulations, as follows: 1) women < 50
and amino acids adequacies (4, 6). In other words, for the years old; 2) women 50–64 years old; 3) women ≥ 65
majority of people in Western countries, who are typically years old; 4) men < 65 years old; and 5) men ≥ 65 years
omnivorous, the risk of inadequate protein intake is low, and old.
it is minimally influenced by the animal protein contribution
to the diet. Yet, caution is warranted for elderly people Dietary data and calculation of mean observed diets
because they have higher protein requirements than other adults Dietary data were derived from the Second French Individual
(estimated at 1 g/kg for adults over 65 y and 0.83 g/kg for and National Study on Food Consumption (INCA2), a 7-
younger adults) (7). Thus, for French adults aged 65 and over, day open-ended food-record representative survey conducted
a higher risk of frailty was observed for protein intakes lower between 2005 and 2007 by the French Agency for Food,
than 1 g/kg of body weight, independent of energy intake levels Environmental, and Occupational Health and Safety (14). For
(8). this study, all adults above the age of 18 y were studied,
Protein-source foods provide several nutrients other than leading to a total sample of 2624 individuals, distributed as
proteins and largely contribute to overall nutrient adequacy 922 women < 50 y, 418 women between 50 and 64 y, 197
(9, 10). Animal-based foods provide nutrients that are either women ≥ 65 y, 936 men < 65 y, and 151 men ≥ 65 y.
not found in plant-based sources (such as vitamins D and B12 The INCA2 survey was approved by the French National
and long-chain omega 3 fatty acids), found in small amounts Commission for Computed Data and Individual Freedom
(e.g., vitamin B6 and riboflavin), or found in less bioavailable (Commission Nationale de l’Informatique et des Libertés,
forms (e.g., iron and zinc), with the latter including unique CNIL).
sources of fiber, folate, vitamins E and C, and other antioxidants. For each of the 5 subpopulations, we estimated the average
In addition, diets with plenty of plant-based foods are not daily dietary intake, also called the observed diet below, based
necessarily more affordable than diets with more animal- on 212 frequently consumed food items (Supplemental Table 1),
based foods (11). More generally, food budget constraints following a previously described matching methodology (15).
are important determinants of food choices, and healthy Based on the French food composition table [CIQUAL 2013
food choices tend to be more expensive than unhealthy ones (16)], a sex-specific nutrient composition database containing
(12). energy, fiber, fats (including linoleic and alpha-linolenic acids;
The goal of this study was to determine the extents to EPA; DHA; and lauric, myristic, and palmitic acids, as well
which decreases in protein intakes, particularly from animal- as total SFA), proteins (including amino acids), carbohydrates,
based sources, could be made without impairing the nutritional 11 vitamins, and 11 minerals was derived for each of the 212
adequacy of the diet and at no additional cost. To reach food items (15). Phytate and amino acid contents in foods were
this goal, mathematical optimization models were developed derived from the International Network of Food Data Systems
to determine the theoretical minimum levels of total dietary databases (15). Average prices for the 212 food items were
proteins, as well as the minimum percentages of animal proteins calculated based on data from the 2006 Kantar Worldpanel (15,
in total proteins that are compatible with the fulfillment of all 17). After removal of alcoholic items from food consumption,
nutrient-based recommendations, without changing the total composition, and price databases, nutrient intakes and costs
energy content and at no additional cost, for 5 subpopulations associated with the consumption of 207 food items were
of French adults differing according to sex and age. estimated for each subpopulation, leading to 5 observed
diets.

Modeling
Material The models developed in this study used a mathematical
Populations of interest optimization technique based on the simplex algorithm. An
Five subpopulations of adults were defined according to nu- optimization model is comprised of variables, constraints, and
trient recommendation levels. First, segmentation was applied an objective function, and the algorithm finds the value that
based on sex because recommended levels of vitamins (A, E, each variable must have to comply with the constraints while
optimizing (i.e., minimizing or maximizing) the value given by
the objective function. In diet optimization models, the variables
MS-Nutrition and MoISA received financial support from the French National are the quantities of foods, and the algorithm finds the quantity
Interprofessional Association of Livestock and Meat (Interbev).
Author disclosures: The authors report no conflicts of interest.
of each food in the modeled diet that is compatible with the
Interbev had no role in the design, implementation, analysis, or interpretation of simultaneous fulfillment of all constraints for the minimum (or
the data. maximum) value of the objective function.
Supplemental Figure 1 and Supplemental Tables 1–4 are available from the In the present study, 2 kinds of models were developed,
“Supplementary data” link in the online posting of the article and from the same
called model set #1 and model set #2. Model set #1 diets
link in the online table of contents at http://jn.nutrition.org.
Address correspondence to FV (e-mail: [email protected]). were developed to determine the minimum protein contents
Abbreviations used: INCA2, Second French Individual and National Study on compatible with the fulfillment of all nutrient recommendations
Food Consumption; NRV, nutrient reference values. (without imposing a minimum amount of total proteins),
Animal protein intake and nutritional adequacy 2515
 TABLE 1 Constraints applied to model set #1 and model set #2, for each subpopulation1

2516
Women Men
Subpopulation <50 y 50–64 y ≥65 y <65 y ≥65 y
Nutritional constraints (applied to model set #1 and model set #2)

Vieux et al.
Energy, kcal/d = Observed (1731.6) = Observed (1711.5) = Observed (1675.4) = Observed (2207.1) = Observed (2082.8)
Carbohydrates, % energy [40; 55] [40; 55] [40; 55] [40; 55] [40; 55]
Fats, % energy [35; 40] [35; 40] [35; 40] [35; 40] [35; 40]
Linoleic acids, % energy ≥4 ≥4 ≥4 ≥4 ≥4
Alpha-linolenic acids, % energy ≥1 ≥1 ≥1 ≥1 ≥1
DHA + EPA, mg/d ≥250 ≥250 ≥250 ≥250 ≥250
SFA, % of energy ≤12 ≤12 ≤12 ≤12 ≤12
Lauric + myristic + palmitic acids, % energy ≤8 ≤8 ≤8 ≤8 ≤8
Total sugars, without lactose, mg/d ≤100 ≤100 ≤100 ≤100 ≤100
Sodium, mg/d ≤ Observed (2613.0) ≤ Observed (2624.7) ≤ Observed (2689.1) ≤ Observed (3459.9) ≤ Observed (3575.5)
Sodium-to-potassium ratio, molar ≤1 ≤1 ≤1 ≤1 ≤1
Water, ml/d 2000 2000 2000 2500 2500
Fibers, g/d ≥30 ≥30 ≥30 ≥30 ≥30
Vitamin A, μg/d [650; 3000] [650; 3000] [650; 3000] [750; 3000] [750; 3000]
Thiamin, mg/MJ ≥0.14 ≥0.10 ≥0.10 ≥0.14 ≥0.10
Vitamin B12, μg/d ≥4 ≥4 ≥4 ≥4 ≥4
Riboflavin, mg/MJ ≥0.17 ≥0.17 ≥0.17 ≥0.17 ≥0.17
Niacin, mg NE/MJ;mg/d [1.6; 900] [1.6; 900] [1.6; 900] [1.6; 900] [1.6; 900]
Pantothenic acid, mg/d ≥4.7 ≥4.7 ≥4.7 ≥5.8 ≥5.8
Vitamin B6, mg/d [1.5; 25] [1.6; 25] [1.6; 25] [1.8; 25] [1.7; 25]
Folates, μg/d ≥330 ≥330 ≥330 ≥330 ≥330
Vitamin C, mg/d ≥110 ≥110 ≥110 ≥110 ≥110
Vitamin D, μg/d [5; 50] [5; 50] [5; 50] [5; 50] [5; 50]
Vitamin E, mg/d [9.9; 300] [9.9; 300] [9.9; 300] [10.5; 300] [10.5; 300]
Calcium, mg/d [960; 2500] [950; 2500] [950; 2500] [960; 2500] [950; 2500]
Copper, mg/d [1.0; 5.0] [1.3; 5.0] [1.3; 5.0] [1.3; 5.0] [1.6; 5.0]
Iron, mg/d ≥162 ≥11 ≥11 ≥11 ≥11
Iodine, μg/d [150; 600] [150; 600] [150; 600] [150; 600] [150; 600]
Magnesium, mg/d ≥360 ≥360 ≥360 ≥420 ≥420
Phosphorus, mg/d ≥700 ≥550 ≥550 ≥700 ≥550
Selenium, μg/d [70; 300] [70; 300] [70; 300] [70; 300] [70; 300]
Zinc, mg/d [7.5; 25] [7.5; 25] [7.5; 25] [9.4; 25] [9.4; 25]
≥0.0058 × phytates + 5.8 ≥0.0058 × phytates + 5.8 ≥0.0058 × phytates + 5.8 ≥0.0077 × phytates + 7.1 ≥0.0077 × phytates + 7.1
Nonnutritional constraints (applied to model set #1 and model set #2)
Diet cost, €/d ≤ Observed (5.16) ≤ Observed (5.77) ≤ Observed (5.52) ≤ Observed (6.33) ≤ Observed (6.30)
Fish, g/d <28.6 <28.6 <28.6 <28.6 <28.6
Quantity per food item,3 g/d ≤ 95th pctl ≤ 95th pctl ≤ 95th pctl ≤ 95th pctl ≤ 95th pctl
Fortified products and mineral waters,4 g/d ≤ Observed ≤ Observed ≤ Observed ≤ Observed ≤ Observed
(Continued)
 minimum quantity of total proteins), without changing the total energy content, at no additional cost, and taking eating habits into account. Values in brackets are ranges e.g., in the case of fats, the recommended amount is between 35 and 40%
without changing the observed energy content and without

Fortified foods were defined as breakfast cereals (3 food items), multivitamin juice, soya-based drink, tomato soup, soya-based desserts (3 food items), cocoa powder, pineapple juice, a mixture of oils, instant drink, and low-fat margarine. Each
Model set #1 focused on determining the theoretical minimum level of total dietary proteins compatible with the fulfillment of all nutrient-based recommendations (without imposing a minimum quantity of total proteins), without changing the
Gradual decrease by 5%
exceeding the observed diet cost. Model set #2 diets were aimed

[5th pctl; 95th pctl]

[5th pctl; 95th pctl]
[5th pctl; 95th pctl]

total energy content and at no additional cost. Model set #2 focused on determining the minimum percentage of animal proteins in total proteins compatible with the fulfillment of all nutrient-based recommendations (without imposing a
at determining the minimum percentages of animal proteins in

≥65 y

steps
total proteins compatible with the fulfillment of all nutrient-
based recommendations without changing the observed energy
content, without exceeding the diet cost, and taking eating
habits into account.
Men

Variables.
Variables were the same for the 2 kinds of models. All 207 food
items in the subpopulation-specific database were the variables
Gradual decrease by 5% for each model performed.
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]
<65 y

steps
Constraints.
Constraints applied to models are described in Table 1.
Nutritional constraints were applied to both model set #1
and model set #2. Each model included a subpopulation-
specific set of nutritional constraints imposing the fulfillment
of all nutrient-based recommendations (except proteins, for
which levels were analyzed as output). The energy contents
of modeled diets were kept equal to those of observed diets
Gradual decrease by 5%

(i.e., modeled diets were isoenergetic with observed diets).

[5th pctl; 95th pctl]
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]

All nutrient reference values (NRV) were taken from the

≥65 y

French Agency for Food, Environmental and Occupational

steps

Health and Safety reports (7, 13) available at the start of the
food item was constrained to remain below its observed amount. Mineral waters (8 food items) were constrained in the same way to favor tap water.

project. EPA + DHA and vitamin D constraints were adapted

because they were too constraining (official recommendations
are 500 mg/d for EPA + DHA and 15 μg/d for vitamin D). The
EPA + DHA intake was set at 250 mg/d, in agreement with the
European Food Safety Authority dietary reference value (18),
and the vitamin D constraint was set at 5 μg/d, because this
Gradual decrease by 5%

intake is considered as adequate by various official authorities

[5th pctl; 95th pctl]
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]

(19). As the recommendations for calcium differ before and

Women
50–64 y

The 5th percentile and 95th percentile were estimated for all individuals, including nonconsumers of the food group.

after the age of 25 y, NRV were weighted according to the

steps

percentages of individuals older and younger than 25 y among

women under the age of 50 and men under the age of 65.
Sodium (no quantified recommendation in France) was required
to be maintained below observed intake levels. In accordance
with the French NRV, the recommended levels of zinc applied
to the modeled diets depended on the levels of phytates in diets.
In accordance with French food-based dietary guidelines, the
content of fish was maintained at less than 200 g/wk to limit
Gradual decrease by 5%
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]
[5th pctl; 95th pctl]

fish-related contaminant exposure.

In both model set #1 and model set #2, an applied constraint
<50 y

Quantities < 95th percentile estimated only for consumers of the food item.
steps

where the diet cost was kept below or equal to the observed diet
cost was applied to all models, in order to consider the realism
Under the assumption of high iron losses through menstrual blood.

and affordability of the modeled diets. A maximum quantity

was applied to all models for each of the 207 food items, to
avoid including an unrealistic amount for a given food item.
Other constraints were only applied to model set #2.
Consumption constraints, applied as minimum and maximum
amounts by food group, subgroup, and categories, were
Animal protein contributions to total proteins, %
Additional constraints applied to model set #2 only

introduced in order to better accommodate current eating

habits in diets obtained with model set #2. Moreover, model
of energy. Abbreviation: pctl, percentile.

set #2 diets were used to find the minimum percentages of

Quantity per food subgroup,5 g/d

animal proteins in total proteins that are compatible with the

Quantity per food category,5 g/d
TABLE 1 (Continued)

Quantity per food group,5 g/d

fulfillment of nutrient-based recommendations. To do so, a

constraint was introduced and progressively strengthened to
reduce, by 5% steps, the percentage of animal proteins in total
proteins, leading to a group of different model set #2 diets
Subpopulation

for each subpopulation. The first model of the set did not
include any constraint on animal protein contributions to total
proteins. Then, starting from the percentage of animal proteins
in total proteins reached in the first model, that percentage was
1

2
3
4

Animal protein intake and nutritional adequacy 2517

progressively reduced by 5% steps to obtain the other model set ways. First, we assessed the respective recommended amounts
#2 diets until no solution could be found. by considering the digestible protein content (quality):
AAi
Objective function. Quali AAi = × 100 (3)
AArecoi × Prot DIG
In model set #1, total proteins (in grams) were minimized by
Here, Quali AAi is the quality indicator of indispensable amino
applying the following objective function:
acid i in the diet, AAi is the diet’s digestible amino acid content

207
i (mg/d), AArecoi is the recommended amount of indispensable
Min Prot = Qopt
i ∗ Proti (1) amino acid i (mg/g of proteins), and Prot DIG is the digestible
i=1 protein content in the diet (g/d).
Here, Prot is the protein quantity in the modeled diet, Qopt i is Second, we assessed the respective recommended amounts
the modeled quantity of food i, and Proti is the protein content in the context of a diet achieving the minimum recommended
(in grams) of food i. amount of total proteins (quantity):
In model set #2, the objective function was designed to AAi
minimize the deviation from the observed diet in order to Quantity AAi = × 100 (4)
AArecoi × Prot reco
maximize acceptance, and was calculated as the sum of absolute
Here, Quantity AAi is the indicator of indispensable amino
differences between the quantity of each food from the 207
acid quantity I in the diet, AAi is the diet’s digestible amino
foods listed in the observed diet and the quantity of each food
acid content i (mg/d), AArecoi is the recommended amount of
selected in the modeled diet, as follows:
indispensable amino acid i (mg/g of proteins), and Prot reco is
207 
  the recommended amount of total proteins (expressed in g/d),
 opt 
Min D = Qi − Qobs
i  (2) with the latter being a function of the average body weight of
i−1 the subpopulation.
Here, D is the absolute departure (in grams) between the
modeled and observed diets, Qopt i is the modeled quantity Analyses
of food i, and Qobs i is the observed quantity of food i Analyses were repeated for each of the 5 subpopulations. First,
(estimated over the whole subpopulation sample, including the minimum total protein content achievable with model
nonconsumers). set #1 (i.e., the lowest theoretical protein level compatible
with the fulfillment of all nutrient-based recommendations,
Selection of 1 model set #2 diet for each except the recommendation on proteins, for which levels were
subpopulation analyzed as output, at no additional cost) was reported and
For each subpopulation, among the model set #2 diets, the diet compared to both the recommended and observed intakes of
with the lowest animal protein contribution to total proteins, proteins, separating results into animal and plant proteins.
compatible with nutrient and protein adequacy, affordability, Then, the protein contents in diets modeled with model set
and eating habits, was selected by keeping the diet that followed #2 (i.e., diets with progressive 5% step reductions in animal
the 3 following nutritional and consumption criteria: protein contributions to total proteins that are compatible
with the fulfillment of all nutrient-based recommendations,
1) Adequate total protein content: the total protein content
except the recommendation on proteins, which was analyzed
must not fall below the minimum level recommended
as output, at no additional cost and taking eating habits into
for that subpopulation (so that the selected diet is fully
account) were reported and compared to recommended levels,
nutritionally adequate because it fulfills all nutrient-
separating results into animal and plant proteins. Finally, for
based recommendations, including the recommendation
each subpopulation, the food content of the model set #2
for proteins);
selected diet (i.e., among the model set #2 diets, the diet with the
2) Limited food mass deviation from the observed diet:
lowest animal protein contribution to total proteins, compatible
the objective function value (i.e., the sum of absolute
with nutrient and protein adequacy, affordability, and eating
differences between the modeled and observed quantities
habits) was compared to the same food content for the observed
of each food) must be lower than the total mass of the
diet.
observed diet (in kilograms); and
3) Limited exclusion of food items in the modeled diet:
the total number of foods does not drop by 15% or
more compared to the previously modeled diet (i.e., the Results
diet modeled with a 5-point higher percentage of animal
Table 2 shows the total and animal protein contents in the mean
proteins in total proteins).
observed diets and in diets obtained with model set #1, for each
subpopulation.
Protein adequacy In the observed diets, regardless of the subpopulation,
It was considered that the diets (observed and modeled) dis- the total protein contents were higher than the minimum
played protein adequacy when: 1) the total protein content was recommended protein intake. By construction, the protein
higher than the recommended amount for the subpopulation; contents in all diets obtained with model set #1 are the minimum
2) the quality of indispensable amino acids (histidine, isoleucine, protein quantities that would theoretically be compatible
leucine, lysine, methionine + cysteine, phenylalanine + tyrosine, with meeting all other nutrient recommendations, for each
threonine, tryptophan, valine) was adequate; and 3) the quantity subpopulation. Regardless of the subpopulation, that minimum
of indispensable amino acids was adequate. protein quantity was more or less equal to 50 g/d, well below
Qualitative and quantitative assessments of amino acid observed intakes (around 70 g/d and 90 g/d in women and men,
contents were performed by comparing their contents with respectively) and also below the recommended protein intake,
recommended amounts (Supplemental Table 2), expressed in 2 except for the subpopulation of women < 50 y.
2518 Vieux et al.
TABLE 2 Recommended protein intakes, total protein contents, and animal protein contents in OBS and in diets obtained with
MOD1, for each subpopulation1

Total proteins Animal proteins

Minimum
Mean body recommended intake In g/d In % energy In g/d In % of total proteins
Subpopulation weight, kg of total proteins,2 g/d OBS MOD1 OBS MOD1 OBS MOD1 OBS MOD1
Women < 50 y 62.6 52.0 69.5 54.3 16.0 12.5 49.3 27.6 71.0 50.8
Women 50–64 y 67.7 56.2 71.5 51.0 16.7 11.9 51.2 27.2 71.6 53.3
Women ≥ 65 y 65.5 65.5 68.2 51.6 16.3 12.3 46.9 30.3 68.7 58.8
Men < 65 y 77.7 64.5 93.9 48.4 17.0 8.8 66.5 22.4 70.8 46.3
Men ≥ 65 y 78.8 78.8 88.0 48.0 17.0 9.2 59.6 24.7 67.8 51.4
1
MOD1 focused on determining the theoretical minimum level of total dietary proteins compatible with the fulfillment of all nutrient-based recommendations (without
imposing a minimum amount of total proteins), without changing the total energy content and at no additional cost. Abbreviations: MOD1, model set #1; OBS, observed diets.
2
The recommended intake was estimated by multiplying the average body weight by the recommended intake, expressed in g/kg of body weight.

In the observed diets, regardless of the subpopulation, the (despite no constraints applied to them), with animal protein
protein contributions to total energy were between 16% and contributions to total proteins as low as 40% (men < 65
17%. In contrast, in the model set #1 diets, the protein y), 45% (women 50–64 y), and 50% (women < 50 y). The
contributions to total energy dropped to between 12.5% (in cost constraint was binding in all the modeled diets (data
women < 50 y old) and 8.8% (in men < 65 y old). In not shown). All diets with total protein contents higher than
the observed diets, the animal protein contributions to total the recommended levels were qualitatively and quantitatively
proteins were around 68% to 72%, and they decreased to adequate in terms of their amino acid contents, so that full
percentages between 46.3% (in men < 65 years old) and 58.8% protein adequacy was ensured (Supplemental Table 4).
(in women ≥ 65 years old) in model set #1 diets. Supplemental Figure 1 shows the departures from the
Supplemental Table 3 shows nutrient contents, costs, and observed diets (i.e., the value, D, of the objective function; that
fish contents in observed diets and model set #1 diets for the is, the sum of the absolute differences between the quantity of
5 subpopulations. Some values were exactly equal to values im- each food in the observed diet and the corresponding modeled
posed by their respective (minimum or maximum) constraints. diet), as well as the numbers of remaining food items, induced
Such constraints are named “binding” or “active” constraints. by all model set #2 diets. Depending on the subpopulations,
Identifying them helps to point out which constraints are more departures between 1.0 kg/d and 1.8 kg/d were needed to obtain
difficult to fulfil than others. Binding constraints are also the the model set #2-NO diets. Then, a progressively decreasing
ones with the greatest influence on the kinds and amounts animal protein contribution to total proteins increased the
of food introduced in modeled diets (i.e., the selections or departure from the observed diet and decreased the number of
“food choices” made by the models). Constraints to energy foods in the modeled diet. The subpopulation’s specificities were
(equality), water (equality), SFA (maximum), fiber (minimum), observed as follows:
zinc (minimum), and vitamin D (minimum) were binding for
all 5 subpopulations, as were constraints to the diet cost 1) For women < 50 y, imposing a 50% animal protein
(maximum) and fish content (maximum). Minimum constraints contribution to total proteins induced a departure (D)
to alpha-linolenic acids and to calcium were binding for almost from the observed diet that was higher than the total mass
all subpopulations. Many other constraints were binding for of the observed diet.
some but not all subpopulations. Thus, the minimum constraint 2) For women 50–64 y, imposing a 45% animal protein
to iodine was binding in all diets modeled for women, but not contribution to total proteins induced a departure (D)
for men. Conversely, constraints to total sugars (maximum) and from the observed diet (2.8 kg/d) that was higher than
magnesium (minimum) were binding in all diets modeled for the total mass of the observed diet (2.5 kg/d).
men, but not for women. The constraint to iron was binding 3) For women ≥ 65 y, the modeled diet with a 50% animal
only in the subpopulation of women < 50 years old. protein contribution to total proteins did not reach the
Figure 1 shows the protein contents of observed diets and recommended protein level.
diets obtained with model set #2. In an initial step, and 4) For men < 65 y, the departure from the observed diet
compared to observed diets, fulfilling all the applied constraints was never higher than the total mass of the observed diet,
increased the total protein content for all subpopulations, and the protein contents in modeled diets were always
whereas no constraint was applied to proteins (model set #2- higher than the recommended level. However, the number
NO diets). Then, imposing a decrease in the animal protein of food items dramatically decreased (−17%) between
contribution to total proteins (other diets in model set #2) the modeled diets with 45% and 40% animal protein
induced a progressive decrease in the total protein content. contributions to total proteins.
In most modeled diets, as shown by the bars exceeding the 5) For men ≥ 65 y, the modeled diet with a 55% animal
horizontal black line, the recommended level of protein intake protein contribution to total proteins did not reach the
was achieved despite the absence of a constraint applied to recommended protein level (squares).
it. However, for older subpopulations, the modeled diets did Based on these results and the criteria described above to
not contain the recommended protein levels when the animal select 1 model set #2 diet per subpopulation, modeled diets
protein contributions to total proteins were ≤55% for men ≥ 65 with the following percentages of animal proteins in total
y and ≤50% for women ≥ 65 y. In other subpopulations, proteins were selected for further analysis (Figure 2): 55% for
the modeled diets fulfilled all nutritional constraints, and their women < 50 y, 50% for women 50–64 y, 55% for women ≥ 65
protein contents were higher than the recommended levels y, 45% for men < 65 y, and 60% for men ≥ 65 y.

Animal protein intake and nutritional adequacy 2519

FIGURE 1 Animal proteins, plant proteins, and total protein contents (g/d) in OBS and in diets obtained with MOD2, MOD2-NO, and a
progressively decreasing constraint (in 5% steps) on the animal protein contribution to total proteins2 , until no solution could be found, in (A)
women < 50 y, (B) women 50–64 y, (C) women ≥ 65 y, (D) men < 65 y, and (E) men ≥ 65 y. MOD2 imposed the fulfillment of all nutrient-based
recommendations, except the recommendation for proteins, while minimizing the departure from the observed diet. MOD2-NO was a MOD2
model without constraints on animal protein contributions to total proteins. As an example of the decreasing constraints, MOD2-55 imposed a
maximum animal protein contribution of 55% of the total proteins. The horizontal black line indicates the protein intake level recommended for
the subpopulation. Abbreviations: MOD2, model set #2; MOD2-NO, without model set #2; OBS, observed diets.

Figure 3 shows the dietary shifts in food groups induced by diets, as did unrefined starchy foods, milk, and eggs (except
each selected model set #2 diet, and Table 3 displays the dietary the latter in men < 65 y). In comparison, intakes of
shifts induced by the selected models in more detail. “Water refined starchy foods, breakfast cereals (except in men < 65
and drinks” and “plant-based alternatives” were removed from y), yogurt (except in men ≥ 65 y), cheese, meat and
the graph for visualization purposes because they presented deli meat, plant-based dishes, biscuits and sugar, desserts,
quantities that were too high and too low, respectively, but animal fats (except in men ≥ 65 y), and spices and
were included in Table 3. In all selected modeled diets, fruits sauces (except in women ≥ 65 y) decreased. Fish intakes
and vegetables, dairy products, and starchy foods increased attained the maximum applied amount (200 g per week).
(except the latter in men ≥ 65 y), while other food groups Tea and coffee intakes remained constant. Other subgroups
decreased. from the water and drinks group had decreased intakes
All subgroups within the fruit and vegetable food group for women and remained constant for men. For other
had increased intakes between the observed and modeled food subgroups (animal-based dishes, cakes and tarts, and

2520 Vieux et al.

 TABLE 3 Group and subgroup quantities in OBS and MOD2 selected diets for each subpopulation1

Women < 50 y Women 50–64 y Women ≥ 65 y Men < 65 y Men ≥ 65 y

Food groups and subgroups, g/d OBS MOD2-55%2 OBS MOD2-50%2 OBS MOD2-55%2 OBS MOD2-45%2 OBS MOD2-60%2
Fruit and vegetables 278.3 617.7 455.8 671.3 483.5 756.6 298.1 699.6 475.1 844.0
Fruits (without juice) 112.5 300.7 202.9 352.7 203.0 351.9 122.5 327.0 200.3 432.9
Nuts and seeds 1.58 2.86 2.64 6.79 2.52 13.33 2.74 2.86 1.69 2.50
Vegetables 164.2 314.1 250.2 311.9 278.1 391.4 172.8 369.8 273.1 408.5
Starchy foods 190.1 263.3 177.8 257.3 199.3 236.3 272.5 361.3 290.9 273.3
Unrefined3 63.40 163.6 62.90 155.71 67.76 164.3 78.49 190.0 86.66 160.6
Refined 121.1 97.10 111.0 99.15 129.3 71.23 189.2 166.4 202.6 111.2
Breakfast cereals4 5.59 2.59 3.86 2.44 2.27 0.78 4.89 4.89 1.71 1.50
Dairy products 189.9 294.7 196.5 245.4 179.1 271.0 202.0 426.8 161.3 388.0
Milk 90.01 220.9 62.56 157.4 79.71 194.8 98.18 369.2 60.05 321.4
Yogurt 76.28 73.76 109.5 88.00 73.64 72.31 68.47 57.61 59.49 59.49
Cheese 23.57 0.00 24.50 0.00 25.74 3.87 35.41 0.00 41.80 7.10
Meat, fish, and eggs 129.3 95.61 145.4 119.9 134.5 101.3 181.3 77.41 177.6 131.4
Meat and deli meat 89.41 23.55 94.87 45.58 84.61 29.27 138.0 45.07 123.9 58.18
Fish 26.84 28.57 35.42 28.57 34.75 28.57 29.75 28.57 34.24 28.57
Eggs 13.05 43.49 15.11 45.71 15.15 43.49 13.56 3.77 19.45 44.64
Mixed dishes 114.9 124.0 82.30 36.41 79.53 70.66 152.1 18.01 116.6 105.3
Animal-based dishes5 89.59 124.0 58.75 36.02 48.23 69.02 123.1 11.37 68.01 57.90
Plant-based dishes6 25.27 0.00 23.55 0.39 31.31 1.64 29.03 6.63 48.58 47.36
Sweet products 105.1 59.43 94.79 46.48 87.96 59.81 115.6 85.20 84.61 78.02
Biscuits and sugar 32.53 12.02 29.52 8.20 24.17 10.60 35.71 0.15 26.79 2.77
Desserts 15.36 10.11 12.30 8.25 17.59 14.71 16.91 6.53 11.55 10.14
Cakes and tarts 57.19 37.30 52.97 30.02 46.20 34.50 62.97 78.52 46.27 65.11
Plant-based alternatives 3.07 2.79 9.38 9.38 6.99 6.99 2.43 2.43 1.49 1.49
Alternatives to meat 0.28 0.00 0.45 0.45 0.00 0.00 0.07 0.07 0.19 0.19
Alternatives to dairy4 2.79 2.79 8.93 8.93 6.99 6.99 2.36 2.36 1.30 1.30
Added fats 39.22 20.14 42.35 18.69 46.11 28.68 40.35 44.02 49.25 48.66
Animal fats 12.00 0.00 11.18 0.00 12.46 0.00 12.76 8.52 13.91 13.91
Vegetable fats 18.95 17.29 24.64 16.94 27.43 15.82 19.06 32.47 28.81 32.31
Spices and sauces 8.26 2.85 6.53 1.75 6.22 12.86 8.53 3.04 6.52 2.44
Waters and drinks 1305 900.6 1336 969.4 1157 836.2 1265 1265 1086 1086
Water 790.4 491.0 804.1 476.0 757.2 459.3 775.1 775.1 685.8 685.8
Sugary drinks 85.51 0.00 20.02 0.00 9.48 0.00 109.2 109.2 15.60 15.60
Fruit juices (100%) 60.88 41.03 48.37 30.18 31.59 17.77 57.39 57.39 41.26 41.26
Tea and coffee 368.5 368.5 463.2 463.2 359.1 359.1 323.3 323.3 343.0 343.0
1
For each subpopulation, the selected diet is the diet, among the MOD2 diets, with the lowest animal protein contribution (in percentage of total proteins) fulfilling all nutrient-based recommendations, including the recommendation on proteins,
without changing the energy content, at no additional cost, and taking eating habits into account. Abbreviations: MOD2, model set #2; OBS, observed diets.
2
Numbers for the MOD2 diets indicate the animal protein percentages of the total protein in the selected modeled diet.
3
Including legumes (lentils, white beans, etc.).

Animal protein intake and nutritional adequacy

4
Quantities of all fortified foods (e.g., breakfast cereals or plant-based alternatives to dairy) were constrained to not increase compared to observed amounts.
5
Including animal-based salted tarts (such as quiches), sandwiches, burgers, couscous, paella, and so forth.
6
Including plant-based salted mixed dishes (such as tabouleh) and appetizers or biscuits.

2521
 to their low average body weight), the strict minimum was
even higher than the recommended protein level. Numerous
nutritional constraints were binding in model set #1, showing
that foods that are a source of protein were needed for several
other nutrients. Achieving the recommended levels was difficult
for vitamin D, zinc, fiber, alpha-linolenic acid, and SFA in all
subpopulations and for iodine, calcium, magnesium, and total
sugars in some subpopulations. Regarding iron, the constraint
was binding only for young women. In that subpopulation, the
level of the iron constraint (i.e., 16 mg/d) imposed corresponded
to a hypothesis of high iron losses in menstrual blood. Note that
alternative models (data not shown) were conducted where only
11 mg/d of iron (recommended iron intake for young women
with normal iron losses) was required, and the minimized total
protein contents did not change (54 g).
Despite the absence of any constraint related to amino
acid contents in diets, indispensable amino acids were sys-
FIGURE 2 Animal and plant protein contributions to total proteins
(in percentages) in the model set #2 selected diets (i.e., modeled tematically adequate in all modeled diets fulfilling nutrient
diets with the lowest animal contributions to total proteins that recommendations, including the recommendation for total
are compatible with nutrient and protein adequacy, affordability, and proteins, whether their animal protein contribution to total
eating habits) for each subpopulation. The models imposed the proteins was decreased or not. This means that other nutrient
fulfillment of all nutrient-based recommendations except for the deficiencies would occur before amino acid deficiency would
recommendation for proteins, while minimizing the departure from become a problem. Accordingly, previous studies showed that
the observed diet. protein adequacy is slightly or not influenced by the amino
acid distributions in foods that are a source of protein (4), and
is more significantly related to protein quantity than quality,
vegetable fats), the directions of variation differed between except in diets containing high plant protein contributions to
subpopulations. total proteins (around 70%) from minimally diversified sources,
such as refined grains (20).
Foods that were sources of both animal and plant proteins
were needed to cover nutrient requirements, even when proteins
Discussion
were minimized. Shifting towards more plant-based diets in
It is known that foods that are sources of protein contain Western countries is often recommended by the scientific
several nutrients other than protein but, to the best of our community and public stakeholders for health and climate
knowledge, this study is the first that assessed to what purposes (1,21–23), but there is no consensus regarding the
extent total proteins and animal protein contributions to total adequate ratio of animal protein to plant protein in sustainable
proteins could be theoretically reduced without impairing diets. A 1:1 ratio (1 g of plant protein for 1 g of animal protein)
the fulfillment of all other nutrient-based recommendations, is often presented as a nutritional standard but, to the best of
excluding the use of nutritional supplements or fortified foods. our knowledge, it is not recommended by any official order. In
A mathematical optimization approach was used to model this study, models decreasing the animal protein contributions
population diets while simultaneously fulfilling a set of nutrient- to total proteins by 5% steps (model set #2) showed that animal
based constraints (not including the constraint on proteins), protein contributions of 60% (men ≥ 65 y), 55% (women < 50
without changing dietary energy and at no additional cost. The y and ≥ 65 y), 50% (women 50–64 y), and 45% (men < 50
results showed that in highly theoretical models where total y) would be fully compatible with adequate nutrients and
proteins were directly minimized without considering eating proteins, affordability, and eating habits, despite their relatively
habits, a strict minimum of at least 48 g/d of total proteins was low protein contents. Our results therefore imply that moving
needed to meet nutrient-based recommendations for nutrients towards diets with lower animal protein contributions or to
other than proteins. Models better at taking eating habits into fully vegan diets, like the ones included within the range
account showed that the animal protein contributions to total of diets recommended by the EAT–Lancet Commission (21),
proteins, which were approximately 70% in observed diets, would necessarily require food fortification and/or nutrient
could be reduced to contributions that were between 45% and supplementation to cover adult nutritional requirements. The
60%, depending on age and sex, while still being compatible minimum percentages of animal proteins found in the present
with complete nutritional adequacy and affordability. Lower study to be compatible with nutrient adequacy in the different
percentages would either be mathematically unattainable or subpopulations considered can be compared to those obtained
constraints would have to be relaxed or removed, therefore in 2 recent studies focused on designing sustainable diets: 1
impairing nutritional adequacy and/or realism in the resulting for French adults from the NutriNet-Santé cohort (24) and
modeled diets. 1 for older Dutch adults from the Longitudinal Aging Study
One important finding from this study is that in the absence Amsterdam cohort (25). In apparent contradiction to our
of any constraint imposing a given protein level, a strict results, for the French study, the mathematical optimization
minimum protein amount was needed to cover other nutrient- approach used to derive nutritionally adequate individual diets
based recommendations in adults (model set #1). This quantity with progressively lower environmental impacts, while also
was close to the minimum recommended protein intake level controlling for costs, resulted in diets with an animal protein
for each subpopulation. For the subpopulation of women < 50 contribution to total protein as low as 22% for the most so-
y (where the recommended protein level was relatively low due called “disruptive” scenario (24). Such a discrepancy can be
2522 Vieux et al.
FIGURE 3 Observed (x-axis) and modeled (y-axis) quantities from selected MOD2 diets of each food group for each subpopulation. For (A)
women < 50 y and (C) women ≥ 65 y, the MOD2 selected diets contained animal proteins as 55% of total proteins; for (B) women 50–64 y, the
MOD2 selected diet contained animal proteins as 50% of total proteins; for (D) men < 65 y, the MOD2 selected diet contained animal proteins
as 45% of total proteins; and for (E) men ≥ 65 y, the MOD2 selected diet contained animal proteins as 60% of total proteins. Abbreviation:
MOD2, model set #2.

Animal protein intake and nutritional adequacy 2523

explained through several methodological differences. Unlike populations, especially children and populations from countries
our study, the NutriNet-Santé study did not consider vitamin with differing protein intake patterns (31).
D and omega 3 fatty acids; fiber was set at a minimum level of In conclusion, this study showed that for this French
only 23 g/d (compared to 30 g/d, the French recommendation, adult population, the lowest animal protein contributions to
in our study); fortified foods appeared to be allowed to increase total proteins that are compatible with nutritional adequacy,
(whereas they were constrained to not increase in our study); affordability, and eating habits vary from 45% to 60%,
the minimum constraint on total proteins was set at only depending on age and sex, with the highest contributions needed
0.66 g/kg of body weight, without any age distinction; and for older populations and young women. The environmental
the amino acid content was not analyzed. As suggested by impacts from resulting dietary shifts need to be assessed in order
the term “disruptive,” a considerable departure from observed to verify their benefits to planetary health.
food consumption patterns was allowed, and 200 g/d of soya-
based products was included (with no information on possible Acknowledgments
fortification). Results from the Dutch study were more in line We thank Florence Garcia-Launay and Joël Aubin, members
with our results, since they showed that it was possible to design of the MINIPROT working group, for their suggestions and
a high-protein (defined as providing > 1.2 g/kg body weight), support.
nutritionally adequate diet with a lower climate impact, a 50% The authors’ responsibilities were as follows – FV, ND:
contribution in animal protein, and lower contributions that designed the research and provided essential materials; FV:
induce amino acid inadequacy (25). This is lower than the 60% conducted the research and analyzed the data; FV, ND,
and 55% contributions we obtained for older men and women, DR: wrote paper; and all authors: reviewed and edited the
respectively, but fewer nutritional constraints were included in manuscript, take responsibility for the final content, and read
the Dutch study than in our models. and approved the final manuscript.
Regardless of the subpopulation, all modeled diets reached
the maximum allowable diet cost (i.e., the observed cost),
confirming that reaching nutritional adequacy tends to be costly
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