CR Biologies

Joseph Igbokwe

Violaine Nicolas

Akinlabi Oyeyiola

Adeoba Obadare

Adetunji Samuel Adesina

Michael Olufemi Awodiran

Natalie Van Houtte

Elisabeth Fichet-Calvet

Erik Verheyen

Ayodeji Olayemi

Molecular taxonomy of Crocidura species (Eulipotyphla: Soricidae) in a key biogeographical region for African shrews, Nigeria

Ouahiba Sihali-Beloui

Djamila Aroune

Fella Benazouz

Adile Hadji

Salima El-Aoufi

Sergio Marco

A hypercaloric diet induces hepatic oxidative stress, infiltration of lymphocytes, and mitochondrial reshuffle in Psammomys obesus, a murine model of insulin resistance

André Nel

252 Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278
Ensuite, aucun serial killer n’est plus efficace: 700 000 morts
par an.
Enfin, sa nuisance ne vient pas de lui, mais des hôtes très per-
nicieux qu’il peut transporter. C’est donc bien un trio qui est à
l’œuvre: un parasite, une femelle moustique qui l’accueille et
l’être humain qu’elle contamine en le piquant pour pomper son
sang. Notons que cette dame n’agit pas ainsi par méchanceté,
mais parce que le sang recueilli contient des protéines néces-
saires au bon développement de ses œufs. En piquant un
humain infecté, elle aspire en même temps le parasite, qui
gagne son estomac, avant, pour notre plus grand malheur, de
gagner ses glandes salivaires: c’est par cette salive qu’il sera
transmis lorsqu’elle piquera un nouvel humain.
La Nature étant, comme l’on sait, sophistiquée, toutes les
espèces de moustiques ne sont pas compétentes pour accueil-
lir tous les virus ou parasites. Par exemple, seuls les Anophèles
transportent Psalmodium falciparum, le responsable du palud-
isme. Et ce sont les Aedes (dont Aedes albopictus, le fameux
« tigre », Fig. 1) qui portent les virus de la dengue, du Zika et du
chikungunya.
Depuis quelques décennies, un double mouvement se dessine.
Le premier est une invasion, l’invasion par nous, les humains,
des habitats animaux. Pour accroître la superficie de nos villes
et de nos champs, nous défrichons, nous avançons dans les
profondeurs des forêts, domaine jusque-là réservé aux bêtes,
petites ou grandes. Il en résulte des épidémies parfois très
graves, comme celles du sida ou de la maladie à virus Ebola.
Car les virus, jadis isolés, sortent de ces zones forestières peu
habitées et se propagent sans frontières.
L’autre mouvement est celui de l’adaptation. Les moustiques
voient en nous de merveilleux repas de sang. Dans ces con-
ditions, pourquoi s’obstiner à piquer la nuit des êtres trop
souvent protégés par des moustiquaires, alors qu’en plein jour
ces mêmes réservoirs de sang se retrouvent sans défense? Et
pourquoi demeurer dans la jungle, alors que les repas de sang
s’accumulent dans les villes? C’est ainsi que les espèces noc-
turnes deviennent diurnes, les forestières, urbaines.
Comment lutter?
D’abord, ainsi que vous le suggère le bon sens, empêcher
les moustiques de se multiplier en supprimant les « gites
larvaires », ces eaux stagnantes où ils naissent et passent
la première partie de leur vie. Ensuite, éviter de se faire
piquer. La prévention, comme chacun sait, est la meilleure
des médecines. Un programme massif de moustiquaires
imprégnées d’insecticide a permis de réduire de moitié le nom-
bre de morts du paludisme.
Des médicaments existent pour contrer les malfaisances du
parasite une fois qu’il nous a infectés. Ainsi, la « poudre des
Jésuites », la future quinine, et l’artémisinine, extraite d’une
plante bien connue de la vieille pharmacopée chinoise. Mal-
gré la combinaison de plusieurs molécules, les phénomènes de
résistance se multiplient, notamment dans l’Est du Cambodge,
une zone où, par chance, la prévalence, c’est-à-dire le nom-
Fig. 1 Moustique tigre - Crédit photo: PHANIE.
bre de personnes atteintes sur une période donnée, reste peu
élevée. De telles résistances en Afrique, plus peuplée et plus
touchée, auraient des conséquences dramatiques.
Ces limites de l’efficacité des médicaments et les difficultés
extrêmes rencontrées pour fabriquer des vaccins ont ouvert
la voie à d’autres méthodes plus radicales. Par exemple, le
lâcher de mâles préalablement rendus stériles dans une zone
de moustiques sauvages. Ou l’utilisation de ciseaux molécu-
laires, le fameux CRISPR-Cas 9, capables de supprimer des gènes
indésirables, mais aussi d’en insérer de nouveaux, comme celui
d’une résistance au paludisme.
Plus fou encore, le forçage génétique, en anglais gene drive, per-
met à une modification génétique réalisée grâce aux ciseaux
moléculaires CRIPR-Cas 9 d’être transmise de façon quasi cer-
taine de génération en génération. C’est ainsi que l’on pourrait
génétiquement programmer l’extinction complète et défini-
tive des moustiques. Certains, rappelant le nombre de décès
causés par ces jolis insectes, y sont déterminés. D’autres sont
beaucoup plus prudents, rappelant leur rôle essentiel dans la
chaîne alimentaire et dans la pollinisation. Ils se demandent
aussi quels « hôtes » les parasites choisiraient si leurs trans-
porteurs habituels disparaissaient. Quant aux manipulations
génétiques, leur sécurité n’est pas encore garantie.
Non scientifique, mais économiste, je voulais continuer mon
travail sur la mondialisation en abordant la santé. Grâce au
docteur de Saint-Aubin, et bien sûr à des dizaines d’experts,
à commencer par mon enseignante en cheffe Anna-Bella Fail-
loux, j’ai pu écrire cette « géopolitique des moustiques ». Qui
confirme, s’il en était besoin, l’unité fondamentale de notre
planète. One health. La vie ne connaît pas de frontières, ni géo-
graphiques, ni entre les règnes.
Déclaration de liens d’intérêts L’auteur déclare qu’il n’a pas de
liens d’intérêts.
https://doi.org/10.1016/j.crvi.2019.09.006
Session II. Diversity, diversification,
extinction
6
The origin of diversity in
insects: Speciation, adaptation
and the Earth dynamics
Philippe Grandcolas
Institut de systématique, évolution, biodiversité (ISYEB),
Muséum national d’histoire naturelle, CNRS, Sorbonne
Université, Paris, France
E-mail address: pg@mnhn.fr
The diversity of insects is often explained as a product of major
radiations, triggered by remarkable adaptations that allowed
them to exploit different environments and to accompany the
rise of flowering plants. A significant part of insect diversity
results however from allopatric speciation with niche con-
servatism that exposed species to different conditions and
stimulated adaptive divergence. We demonstrated that for sev-
eral groups of Insects (cockroaches, crickets and grasshoppers)
in the island of New Caledonia.
Since Darwin and Wallace, islands are considered as labo-
ratories of evolution, being closed systems of manageable
size. New Caledonia is especially interesting in this respect,
according to unique characteristics. Being the oldest oceanic
island in the world and quite isolated from continents, bio-
logical evolution occurred there in situ during long periods.
As a tropical and medium-sized island harboring a very rich
biodiversity, it also offered facilities to study many different
evolutionary questions. New Caledonia (hereafter NC) has thus
the best characteristics as a model system, allowing testing
very diverse evolutionary assumptions with limited efforts
Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278 253
and infrastructures. Unfortunately, these amazing geograph-
ical characteristics have also constrained the scientific study
of NC until now, especially because of its isolation from large
academic centers.
Establishing NC as a modern oceanic model system resulted
however from our work, later than the 1990s. Before, NC had
the reputation of an amazingly old Gondwanan place and was
intuitively considered as a continental fragment. If so, its use-
fulness as a model system to study evolution would have been
limited since the biota of continental islands (e.g., Madagascar
or New Zealand) are a complicated mix of organisms dating
back to the separation with neighboring continental territories
or having colonized the island by dispersal. In most of these
cases, it happened difficult to distinguish between both these
different components and to examine evolutionary assump-
tions that require a reference dating point. The reasons why NC
was traditionally considered a continental territory were the
age of its deep geological basement and the local occurrence
of relict species. In agreement with this common assump-
tion, the NC geological deep basement is ancient and predates
Gondwanan breakup. But this basement has been submitted to
important environmental disturbances because of the island
location at the limit between two tectonic plates. Actually, the
island has been submerged twice for a long time at Paleocene
and Eocene epochs and only emerged around 37 ± 3 Ma.
This geological background was in need of independent test-
ing with biological studies. We thus built multiple molecular
phylogenetic trees of different groups of organisms, dated with
probabilistic methods and external calibration points. A first
review [1] and a more recent meta-analysis [2] of these stud-
ies showed that most groups colonized NC and diversified just
after 37 My, confirming the geological scenario of recent terres-
trial emergence. However, a few local species belong to groups
that were dated much older than 37 My, representing true
relicts, i.e. recent species that remained from old clades [3].
These old clades can only be assumed to have been ancestrally
present in the region, colonizing NC after its emergence and
going extinct in other territories, either drowned islands or
mutated continental ecosystems.
These different results strongly indicated that NC is an oceanic
island, even if very old–actually the oldest one in the world–and
that its local biota dated back to island emergence. Relicts are
fascinating species, unfortunately too sparsely known after the
major extinction of their relatives to support any robust biogeo-
graphic scenario. Fossil data are absolutely needed to assess the
evolutionary history of these taxa without speculation.
Setting up this biogeographic scenery has allowed one to
properly conduct speciation studies by inferring the age and
the ancestral phenotypic characters of local lineages. We did
that in several groups of insects, cockroaches, crickets, and
grasshoppers, and we showed that speciation occurred mainly
in relation to orography, in allopatry and with niche conser-
vatism. Closely related species occurred as narrow endemics
in the same habitats of neighboring areas, mainly on adjacent
small mountains. Sympatry, when observed, was inferred to
be secondarily caused by the increase of distribution area of
older species. Even if most speciation events seemed to have
occurred without any important evolutionary change, some
adaptive shifts have been detected in several cases, from dif-
ferent food plants on metalliferous soils to different habitats in
the forest understory. The major conclusion that can be drawn
from high speciation rate and narrow endemicity in New Cale-
donia is that speciation went before, not always together with
adaptive divergence. Here we offer both a new and powerful
natural laboratory of evolution, calibrated in space and time,
and a model of speciation where niche conservatism is the
engine for adaptive divergence.
Disclosure of interest The author declares that he has no com-
peting interest.
References
[1] P. Grandcolas, J. Murienne, T. Robillard, L. Desutter-
Grandcolas, H. Jourdan, E. Guilbert, et al., New Caledonia:
a very old Darwinian island? Philos. Trans. R. Soc. Lond. B
Biol. Sci. 363 (2008) 3309–3317.
[2] R. Nattier, R. Pellens, T. Robillard, H. Jourdan, F. Legendre, M.
Caesar, et al., Updating the Phylogenetic Dating of New Cale-
donian Biodiversity with a Meta-analysis of the Available
Evidence, Sci. Rep. 7 (2017) 3705.
[3] P. Grandcolas, R. Nattier, S.A. Trewick, Relict species: a relict
concept? Trends Ecol. Evol. 29 (2014) 655–663.
https://doi.org/10.1016/j.crvi.2019.09.007
7
A glance at the deep past history
of insects
André Nel
Institut de systématique, évolution, biodiversité, ISYEB, UMR
7205, CNRS, MNHN, UPMC, EPHE, Muséum national d’histoire
naturelle, Université des Antilles, Sorbonne Université, CP 50,
Entomologie, 57, rue Cuvier, 75005 Paris, France
E-mail address: andre.nel@mnhn.fr, anel@mnhn.fr
With more than 1,100,000 described species, the insects are
the most diverse clade of extant animals, far before all other
groups. Nevertheless, they undergo a drastic decrease of their
populations, due to the sixth extinction of human origin. Thus
it is important to define when and how they became so diverse
and if they were impacted by the major crises of biodiversity
in the deep past to estimate the importance of the current one.
Insects are generally among the best preserved terrestrial fos-
sil organisms, much more complete than the vertebrates. They
are also much more frequent. Thousands can be found in
Konservat-Lagerstätten since the Carboniferous. They are pre-
served either in lacustrine sediments as compression fossils, or
embedded in amber (fossil resins) (Fig. 1).
The Hexapoda (or six-legged arthropods, viz., wingless Collem-
bola, Diplura, Protura ; wingless and winged Insecta) are among
the oldest known terrestrial organisms, with first records dated
from the Middle Devonian of Rhynie in Scotland. Recent molec-
ular phylogenetic dating suggests that they appeared during
the Silurian or even the Ordovician, with the first terrestrial
plants. The Devonian hexapodan record is very scarce and dis-
appointing, with less than six described fossils, all wingless [1].
Fig. 1 A. Compression fossil, Orthoptera, Middle Permian
(China). B. Amber inclusion, Hymenoptera Bethylidae, earliest
Eocene (France). Scale bars: 10 mm (A), 1 mm (B).
A. Fossile en compression, Orthoptera, Permien moyen (Chine).
B. Fossile dans de l’ambre, Hymenoptera Bethylidae, Éocène basal
(France). Échelles: 10 mm (A), 1 mm (B).


A glance at the deep past history of insects

Mariailaria Verderame

Rosaria Scudiero

Health status of the lizard Podarcis siculus (Rafinesque-Schmaltz, 1810) subject to different anthropogenic pressures

Marcel Dicke

Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278 275
Trypanosoma brucei for Tsetse Flies, PLoS. Pathog. 12 (2016)
e1005744.
[5] H.A.T.R. Informal Expert Group on Gambiense, P. Buscher,
J.M. Bart, M. Boelaert, B. Bucheton, G. Cecchi, et al., Do
Cryptic Reservoirs Threaten Gambiense-Sleeping Sickness
Elimination? Trends. Parasitol. 34 (2018) 197–207.
[6] S. Shaw, S.F. DeMarco, R. Rehmann, T. Wenzler, F. Florini, I.
Roditi, et al., Flagellar cAMP signaling controls trypanosome
progression through host tissues, Nat. Commun. 10 (2019)
803.
https://doi.org/10.1016/j.crvi.2019.09.026
Session VI. Insects in the future
26
Insects on the menu
Marcel Dicke
Wageningen University & Research, The Netherlands
E-mail address: marcel.dicke@wur.nl
Major challenges facing humanity Feeding the rapidly
growing human population is a challenge with many facets:
the need for producing 70% more food in 2050 compared to
2010 is challenging in itself. Yet, it is even more challenging
in the context of environmental and population changes,
nutrition crises, and health issues. Moreover, this challenge
is closely connected to other challenges, including mitigation
of climate change, avoiding waste by developing a circular
economy, conserving biodiversity, and supporting human
health (Fig. 1). These challenges can only be met when we
fundamentally change the way we produce food because
current food production methods face serious limitations in
resources, water, land, and energy [1]. The most important
issue in supplying food to the rapidly growing human popu-
lation is the production of sufficient high-quality proteins. At
present, this especially relies on the production of livestock.
However, livestock production has a large ecological footprint
Fig. 1 The challenge of feeding the rapidly growing human
population is closely connected to other challenges, including
mitigation of climate change, avoiding waste by developing
a circular economy, conserving biodiversity, and supporting
human health.
in terms of land use and greenhouse gas production [2,3].
Most of this is the result of the inefficient conversion of
feed-to-meat by conventional livestock, especially cattle
[3]. At present, 80% of global agricultural land is used to
produce livestock, and increasing the production of meat will
have major negative consequences for land use and climate
change. Hedenus et al. [2] concluded that a reduction in
ruminant meat consumption is indispensable for meeting
the maximum 2 ◦C target above the pre-industrial level. An
excellent, sustainable, alternative to conventional meat is
available in insects. Insects are the most abundant group of
animals on Earth with one million species being described and
an estimated 5 million species still to be discovered. More than
2000 insect species are known to be consumed by humans
(https://www.wur.nl/en/Research-Results/Chair-groups/
Plant-Sciences/Laboratory-of-Entomology/Edible-insects/
Worldwide-species-list.htm) (Fig. 2). Their nutritional content
varies between species, but in general the protein content is
similar to that of conventional meat, while insects contain
more unsaturated fatty acids [4]. Moreover, the high mineral
content of insects in comparison to conventional meat is
particularly interesting, considering the worldwide preva-
lence of iron and zinc deficiency. For instance, anaemia is a
global public health problem affecting a quarter of the human
population [5].
Insects in a circular economy Insects can transform rest
streams such as food waste or rest streams from food indus-
try into high-value protein products. In doing so, they are
valuable components of a circular economy. Recent reports
show that insects can make important contributions to global
food security [6] and that producing insects for food and feed
has prospects for rapid commercial and societal uptake. Insect
production has a much smaller ecological footprint, in terms
of land and water use and greenhouse warming potential
compared to the production of pigs and cattle [6–8]. This is
especially due to the much better feed-to-meat conversion
ratio of insects: ca. 2.2 kg feed required per kg of edible weight
production for crickets, whereas this is more than 10 times
higher (25 kg per kg edible weight produced) for beef [4,9].
Moreover, important feed sources for livestock production cur-
rently include fish meal and soybean meal, both obtained from
a market competing between human food and animal feed.
In addition, the use of fishmeal poses an increasing threat to
the viability of marine and aquatic ecosystems [10]. In recent
years, important developments have been initiated in the pri-
vate sector that can contribute to the urgently needed changes
in food production systems, i.e. the production of insects as
food and feed [6,11,12]. Important developments are being
made in the commercial production of insects as food and
feed (www.ipiff.org). These new production systems provide
an important contribution to addressing several of the main
challenges that we face, including resource, land, and water
scarcity [12].
Fig. 2 Menu of a restaurant in Hanoi, Vietnam.
276 Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278
Future To effectively develop insects as food and feed, a first
step should be to rehabilitate insects and leave the ‘nasty,
dirty and avoidable’ image behind. This will require exten-
sive information on the environmental and nutritional benefits
as well as in assuring food safety aspects [13]. In the con-
text of the positive change in the perception of insects as
human food that we have seen in the western world in the
past two decades, it is clear that a lot can be accomplished
in the years ahead. Yet, much research needs to be done for
this, especially in a multidisciplinary and transdisciplinary con-
text. Topics to investigate include, e.g., production of insects on
various rest stream, consequences for insect health and immu-
nity, insects and food quality, economics, consumer behaviour,
food technology, human health aspects, ethics and value chain
development. Such studies will help to develop a new food sec-
tor that is likely to provide the growing human population with
a novel protein source whose production is much more sustain-
able than current production of animal protein. Moreover, this
new food sector also has excellent opportunities to contribute
to achieving various of the global development goals [14,15].
Disclosure of interest The author has not supplied his decla-
ration of competing interest.
References
[1] H.C.J. Godfray, J.R. Beddington, I.R. Crute, L. Haddad, D.
Lawrence, J.F. Muir, et al., Food security: The challenge of
feeding 9 billion people, Science 327 (2010) 812–818.
[2] F. Hedenus, S. Wirsenius, D.J.A. Johansson, The importance
of reduced meat and dairy consumption for meeting strin-
gent climate change targets, Clima. Change 124 (2014)
79–91.
[3] O. Hefferman, Sustainability: A meaty issue, Nature 544
(2017) S18–S20.
[4] A. van Huis, Potential of insects as food and feed in assuring
food security, Annu. Rev. Entomol. 58 (2013) 563–583.
[5] E. McLean, M. Cogswell, I. Egli, D. Wojdyla, B. de Benoist,
Worldwide prevalence of anaemia, WHO Vitamin and
Mineral Nutrition Information System, 1993–2005, Public
Health Nutr. 12 (2009) 444–454.
[6] A. van Huis, J.H.K. Van Itterbeeck, E. Mertens, A. Halloran,
G. Muir, P. Vantomme, Edible insects; Future prospects for
food and feed security, Food and Agriculture Organisation,
United Nations, 2013.
[7] A. Halloran, N. Roos, J. Eilenberg, A. Cerutti, S. Bruun,
Life cycle assessment of edible insects for food protein:
a review, Agron. Sustain. Dev. 36 (57) (2016) 13.
[8] D.G.A.B. Oonincx, I.J.M. de Boer, Environmental impact
of the production of mealworms as a protein source for
humans - A life cycle assessment, Plos One 7 (2012).
[9] A. van Huis, M. Dicke, H. van Gurp, The Insect
Cookbook–Food for a Sustainable Planet, Columbia Uni-
versity Press, New York, 2014.
[10] L. Deutsch, S. Gräslund, C. Folke, M. Troell, M. Huitric, N.
Kautsky, et al., Feeding aquaculture growth through glob-
alization: Exploitation of marine ecosystems for fishmeal,
Global Environmental Change 17 (2007) 238–249.
[11] Y. Hanboonsong, T. Jamjanya, P.B. Durst, Six-legged live-
stock: edible insect farming, collecting and marketing in
Thailand, Food and Agriculture Organisation, Bangkok,
2013.
[12] A. Van Huis, J.K. Tomberlin (Eds.), Insects as food and feed.
From production to consumption, Wageningen Academic
Publishers, Wageningen, The Netherlands, 2017.
[13] M. Dicke, Ecosystem services of insects, in: A. Van Huis, J.K.
Tomberlin (Eds.), Insects as Food and Feed: From Produc-
tion to Consumption., Wageningen Academic Publishers,
Wageningen, The Netherlands, 2017, pp. 60–76.
[14] S.Y. Chia, C.M. Tanga, J.J.A. van Loon, M. Dicke, Insects
for sustainable animal feed: inclusive business models
involving smallholder farmers, Curr. Opin. Environ. Sus-
tain. (2019) [in press].
[15] M. Dicke, Insects as feed and the Sustainable Development
Goals, J. Insects Food Feed 4 (2018) 147–156.
https://doi.org/10.1016/j.crvi.2019.09.027
27
Industrial insect production as
an alternative source of animal
protein
Antoine Hubert
Ÿnsect, 1, rue Pierre-Fontaine, 91000 Évry, France
E-mail address: antoine.hubert@ynsect.com
By 2050, the World Resources Institute projects a 70% human
food calorie gap [1]. The race for sustainable protein alter-
natives is heating up and to avoid global issues, the world
needs massive productivity increases in protein production.
The FAO expects insect protein could help close the gap [2,3].
The European insect sector (Fig. 1) is an emerging industry,
which concentrates on research and innovation efforts that are
invested in the sustainable protein sector worldwide. Legisla-
tive decisions taken by EU policy makers constituted decisive
factors that contributed to boost the advancement of the sector.
We urgently need to address the entire food chain and make it
possible to produce a better, high-quality product–with lower
cost inputs. This means zeroing in on earlier stages of the
food production process: namely how farmed fish and, ulti-
mately, poultry and cattle are fed. Livestock consume roughly
20% of global proteins and compete with humans for ocean
fish stocks, water, land, and soil resources (FAO, 2016). Poul-
try, pigs, aquaculture and some pets are fed diets that include
fishmeal and fish oils, which are produced from ocean-caught
fish and fish trimmings. Aquaculture plays a critical role in
human nutrition, growing faster than any other protein source
for human consumption. Around half of the fish we eat today
comes from farmed sources. Yet fishmeal, the primary food
source for farmed fish, is in crisis because it is derived from
fast-depleting ocean fish stocks.
By replacing traditional animal and fish-based diets with insect
protein, we can offset the growing competition for the ocean
fish stock required to be able to feed 2 billion more people by
2050. This further serves to reduce fish, water, and soil deple-
tion, as well as agriculture’s 25% share of global greenhouse gas
emissions. Insects are part of a natural diet for fish, birds and
some mammals; they contain high levels of essential proteins
and nutrients, which are optimal for animal growth. Today, the
feed given to farmed animals does not include this sustainable
protein.
However, due to the rising costs of traditional feed ingredi-
ents, coupled with the increased production of farmed fish,
feed nutritionists and manufacturers have reduced the amount
of fishmeal in aquaculture diets, replacing it with plant-based
ingredients, especially soy. Mostly imported to Europe from
the Americas, soy bean production carries a huge environmen-
tal cost resulting in, among other things, deforestation and soil
erosion. It is pretty crazy to be feeding the animals we eat with
one of the most expensive forms of protein in the world. This
Fig. 1


Insects on the menu

Antoine Hubert

Industrial insect production as an alternative source of animal protein

Joan van Baaren

276 Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278
Future To effectively develop insects as food and feed, a first
step should be to rehabilitate insects and leave the ‘nasty,
dirty and avoidable’ image behind. This will require exten-
sive information on the environmental and nutritional benefits
as well as in assuring food safety aspects [13]. In the con-
text of the positive change in the perception of insects as
human food that we have seen in the western world in the
past two decades, it is clear that a lot can be accomplished
in the years ahead. Yet, much research needs to be done for
this, especially in a multidisciplinary and transdisciplinary con-
text. Topics to investigate include, e.g., production of insects on
various rest stream, consequences for insect health and immu-
nity, insects and food quality, economics, consumer behaviour,
food technology, human health aspects, ethics and value chain
development. Such studies will help to develop a new food sec-
tor that is likely to provide the growing human population with
a novel protein source whose production is much more sustain-
able than current production of animal protein. Moreover, this
new food sector also has excellent opportunities to contribute
to achieving various of the global development goals [14,15].
Disclosure of interest The author has not supplied his decla-
ration of competing interest.
References
[1] H.C.J. Godfray, J.R. Beddington, I.R. Crute, L. Haddad, D.
Lawrence, J.F. Muir, et al., Food security: The challenge of
feeding 9 billion people, Science 327 (2010) 812–818.
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https://doi.org/10.1016/j.crvi.2019.09.027
27
Industrial insect production as
an alternative source of animal
protein
Antoine Hubert
Ÿnsect, 1, rue Pierre-Fontaine, 91000 Évry, France
E-mail address: antoine.hubert@ynsect.com
By 2050, the World Resources Institute projects a 70% human
food calorie gap [1]. The race for sustainable protein alter-
natives is heating up and to avoid global issues, the world
needs massive productivity increases in protein production.
The FAO expects insect protein could help close the gap [2,3].
The European insect sector (Fig. 1) is an emerging industry,
which concentrates on research and innovation efforts that are
invested in the sustainable protein sector worldwide. Legisla-
tive decisions taken by EU policy makers constituted decisive
factors that contributed to boost the advancement of the sector.
We urgently need to address the entire food chain and make it
possible to produce a better, high-quality product–with lower
cost inputs. This means zeroing in on earlier stages of the
food production process: namely how farmed fish and, ulti-
mately, poultry and cattle are fed. Livestock consume roughly
20% of global proteins and compete with humans for ocean
fish stocks, water, land, and soil resources (FAO, 2016). Poul-
try, pigs, aquaculture and some pets are fed diets that include
fishmeal and fish oils, which are produced from ocean-caught
fish and fish trimmings. Aquaculture plays a critical role in
human nutrition, growing faster than any other protein source
for human consumption. Around half of the fish we eat today
comes from farmed sources. Yet fishmeal, the primary food
source for farmed fish, is in crisis because it is derived from
fast-depleting ocean fish stocks.
By replacing traditional animal and fish-based diets with insect
protein, we can offset the growing competition for the ocean
fish stock required to be able to feed 2 billion more people by
2050. This further serves to reduce fish, water, and soil deple-
tion, as well as agriculture’s 25% share of global greenhouse gas
emissions. Insects are part of a natural diet for fish, birds and
some mammals; they contain high levels of essential proteins
and nutrients, which are optimal for animal growth. Today, the
feed given to farmed animals does not include this sustainable
protein.
However, due to the rising costs of traditional feed ingredi-
ents, coupled with the increased production of farmed fish,
feed nutritionists and manufacturers have reduced the amount
of fishmeal in aquaculture diets, replacing it with plant-based
ingredients, especially soy. Mostly imported to Europe from
the Americas, soy bean production carries a huge environmen-
tal cost resulting in, among other things, deforestation and soil
erosion. It is pretty crazy to be feeding the animals we eat with
one of the most expensive forms of protein in the world. This
Fig. 1
Insects: Friends, foes, and models/Insectes : amis, ennemis et modèles / C. R. Biologies 342 (2020) 247–278 277
is why we need to focus on the start of the food chain. Insects
could make up the protein shortfall without requiring us to
acquire dramatic new tastes in food.
Insects can be a sustainable alternative to soy and fishmeal in
particular. Not only are insects already part of the natural diet
of cattle, pigs, poultry, and fish, but insects deliver an important
source of high-quality protein and polyunsaturated fats.
Farming insects is also hugely cost-efficient and suitable for
many climates. I first began breeding Molitor/mealworm larvae
at Ÿnsect, the company I co-founded in 2011, and discovered
just how energy efficient it was. Insects require very little
water, can be grown in the dark all year round, have no impact
on sea or freshwater biodiversity, and even lead to improve-
ments in land use productivity.
Specialists and scientists have known about the Molitor species
for a long time. The mealworm (the larval form of the Tene-
brio Molitor beetle) is comprised of more than 70% protein
and is a natural source of nutrients for a wide array of animals,
including fish, poultry, pigs, dogs, and cats.
It is also the insect species most suited to industrial devel-
opment. It is a naturally gregarious, nocturnal species, which,
from a practical standpoint, makes breeding easier. It brings
real added value to the market for alternative protein sources
thanks to its nutrient-packed content and unique health ben-
efits for plants and animals.
Our chosen technology and insect species allow us to fully focus
on sustainable development: to do more with less; to use fewer
resources to feed plants and animals; to use fewer antibiotics
and pesticides; to reduce our physical footprint.
Given its intrinsic qualities, the Molitor is perfectly placed to
promote a circular economy. The mealworm consumes all sorts
of organic matter, even low-grade materials. It grows quickly
and requires less space, less earth and less water than other ani-
mal protein sources. It also releases less ammonia and fewer
greenhouses gases than other premium animal proteins. The
entire production system is modelled on a circular economy
with zero-waste. The Molitor protein is produced using fewer
antibiotics and chemical fertilizers and the final product is 72%
protein, with proven nutritional and health benefits for aqua-
culture and pets. With two billion more people to feed within
a matter of 30 years and fish being the fastest-growing source
of protein for human consumption globally, insect protein can
play a pivotal role in closing the gap.
I envision a time when insect protein is at the start of the
food chain for almost everything consumers eat. And even
our plants will be supercharged with fertilizers derived from
insects applied to increase yields.
Putting insects back to their rightful place at the start of the
food chain will give our planet much more breathing space
than misguidedly trying to force everyone to give up meat and
dairy. With insect farming, we are creating a new and highly
sustainable agrifood industry, backed by extremely innovative
tried-and-tested production facilities. Hopefully, the big brand
beasts at the top of the global food chain will soon realize this,
and will back the logical solution to the protein crisis rather
than simply jumping on the popular bandwagon.
Disclosure of interest The author declares that he has no com-
peting interest.
References
[1] J. Ranganathan, D. Vennard, R. Waite, B. Lipinski, T.
Searchinger, P. Dumas, et al., Shifting diets for a
sustainable food future. Creating a Sustainable Food
Future, Installment Eleven, report, World Resources Insti-
tute, 2016 [https://wriorg.s3.amazonaws.com/s3fs-public/
Shifting Diets for a Sustainable Food Future 1.pdf].
[2] Food and Agriculture Organization (FAO), How to feed
the world in 2050, report, 2009, http://www.fao.org/
fileadmin/templates/wsfs/docs/expert paper/How to
Feed the World in 2050.pdf.
[3] P. Rema, S. Subramanian, B. Armenjon, C. Motte, J.
Dias, Graded Incorporation of Defatted Yellow Meal-
worm (Tenebrio molitor) in Rainbow Trout (Oncorhynchus
mykiss) Diet Improves Growth Performance and Nutrient
Retention, Animals 9 (4) (2019) 187, http://dx.doi.org/10.
3390/ani9040187.
https://doi.org/10.1016/j.crvi.2019.09.028
28
The impact of climate on the
winter strategies of insects
Joan van Baaren
UMR-CNRS 6553 ECOBIO, Rennes University, France
E-mail address: joan.van-baaren@univ-rennes1.fr
Life-history traits within ecological communities can be
influenced by two opposite pressures, the first one being
community-wide density-dependent processes like competi-
tion (internal filters), and the second regional environmental
conditions (external filters). While species belonging to a guild
may present contrasting traits as a means of niche differ-
entiation, allowing coexistence whilst exploiting the same
resources, traits in a regional context may converge to a nar-
row range of values because of environmental filtering [1]. As
ectotherm organisms depend strongly on temperatures, con-
vergent winter strategies could be expected in insects of the
same area. In temperate areas, insects can escape the stress-
ful winter conditions either by (i) migration, (ii) diapause
(i.e. arrest of development), or (iii) remaining in activity by
producing cryoprotectant molecules. Concerning the diapause
strategy, it was shown that temperature is both a selec-
tive pressure and a modulator of the diapause expression in
insects. Thus, with climate warming, and especially winter
warming, which acts at the community level, a convergent
alteration of the response to seasonal changes is expected for
the ectotherms, either through genetic adaptations to novel
climatic conditions or through phenotypic plasticity.
In a first study [2], we observed the loss of diapause strat-
egy in two species of an aphid-parasitoid community. Since
the 1980s in western France, the winter guild of aphid par-
asitoids (Hymenoptera: Braconidae) in cereal fields has been
made up of two species: Aphidius rhopalosiphi and Aphidius
matricariae. The recent activity of two other species, Aphidius
avenae and Aphidius ervi, during the winter months suggests
that a modification of aphid parasitoid overwintering strategies
has taken place within the guild. In this study, we first per-
formed a field survey in the winter of 2014/2015 to assess the
levels of parasitoid diapause incidence in agrosystems. Then,
we compared the capacity of the four parasitoid species to
enter winter diapause under nine different photoperiods and
temperature conditions in the laboratory. As predicted, histor-
ically winter-active species (A. rhopalosiphi and A. matricariae)
never entered diapause, whereas the species more recently
active during winter (A. avenae and A. ervi) did enter diapause,
but at a low proportion (maximum of 13.4 and 11.2%, respec-
tively). These results suggest rapid shifts over the last three
decades in the overwintering strategies of aphid parasitoids in
Western France, probably due to winter warming. This implies
that diapause can be replaced by active adult overwintering.
The mechanism behind the change of overwintering strategy
seems to be more a decrease in responsiveness to environmen-
tal signals rather than a genetic loss of diapause. This plasticity
allows non-diapausing individuals to increase their progeny
production. A meta-analysis dedicated to animals has shown
that most of the responses to environmental changes involved
phenotypic plasticity rather than genetic evolution. Moreover,
when genetic changes or shifts in demography, distribution or
phenology occurred, these were generally preceded by a mod-
ification involving phenotypic plasticity. Thus, behavioural
plasticity appears important in explaining the variation in the
success of species to resist to environmental changes.
We confirmed this evolution of winter strategy in our
aphid–parasitoid system, by using a nine-year dataset to build
trophic webs. We confirmed that the community structure
and composition that prevailed before 2011 have recently
shifted toward a more diversified community, with the pres-
ence of the two new braconid parasitoid species studied before


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