THE USE OF VOLATILE ORGANIC COMPOUNDS TO TACKLE

THE SUSTAINABLE DEVELOPMENT GOALS

Iriondo Delgado E., Martín Solís A., Menéndez Linacero C., Rodríguez Serrano M.

ABSTRACT

Plants have complex means of communication. They have shown mechanisms based on
molecules such as neurotransmitters or auxins. However, recent studies have suggested
the importance and relevance of VOCs in plant communication. Here we carried out
bibliographic research on the roles of VOCs and several statistics with worldwide data.

VOCs act enhancing the immunity of the plant itself or those proximate to it, causing an
antioxidant response or inhibiting competitor’s growth. They can be produced
endogenously or by mutualistic microorganisms, the latter being involved in protection
against pathogens. VOC production by biological control agents (BCAs) can be used to
mitigate the harm caused by biotic stresses, generating memory and a more effective
response towards them. Among the world of possibilities they open, one application of
VOCs already put in practice is the push and pull system, designed to reduce input
dependency. For these reasons, VOCs could be an alternative -completely in line with the
sustainable development goals- to pesticides, which entail sustainability problems due to
the contamination and inequality between countries that they aggravate. Nonetheless,
deeper investigation is needed for a wide implementation in agriculture.

INTRODUCTION

It has always been taught that the nervous system is not something that can be applied to
plants. Nevertheless, numerous studies point out there could be similar structures in
plants.

Focusing on the signalling process, the molecule that enables the nervous impulses are
the neurotransmitters, whose mechanism has been found working in plants too (Baluška,
2008). When a plant suffers some external aggression, molecules are released to the rest
of the plan to inform about what is happening, this molecule is the systemin (Ryan, 2000).
Systemin can depolarize the plasma membrane of the cell. Furthermore, glutamate, which
is commonly known to be an important animal neurotransmitter, is apparently a very
abundant molecule in plants. In addition, glutamate receptors in plants are found to be the
same as those in animals.

Another important molecule are auxins, which are hormones that have vital functions in
the plant. It has a very similar way to conduct information in plants as they are transported
within them by vesicles (Baluška et al., 2003).

Something that could have an important role in plants communication are electrical
pulses. A study in a farm in the USA showed evidence that plants can create these
electrical signals that help them to adjust to the environment. Electrodes were placed on
the plant leaves to see how they would react to an insect attack. The farm was suffering
an insect infestation, so it was a perfect stage to perform the experiment. As soon as the
insects were in touch with the plant, the electrodes changed their voltage and it was, not
only transmitted to other parts of the own plant, but to other plants that were around,
(McGowan, 2013) probably to warn them about the threat. It is clear to say that there
must be a way of interaction quite complex that plants use. Therefore, electrical long-
distance signals in plants have been winning popularity among scientists in favour of a
chemical diffusion.

However, one of the principal means of communication plants present is based on volatile
organic compounds (VOCs). These molecules are predominantly terpenes and can be
produced by many forms of life such as plants themselves, insects, fungi, or
microorganisms.

In the case of plants, VOCs can be released in a constitutive manner, or can be induced
by a variety of triggers, including herbivore feeding, insect-egg deposition, a pathogen
attack, or different abiotic stresses. As evidenced by the myriad of situations they are
involved in, VOCs can confer countless distinct signals. This is due to the high structural
diversity of terpenes and the species and organ-specific emission patterns.

Terpene synthases are highly diverse and can produce different alternative skeletons
which in turn can suffer a plethora of post-translational modifications (phosphorylations,
methylations, glycosylations, etc.). All these possibilities enhance the number of terpene
chemical structures plants can emit, allowing them to produce species-specific terpene
blends. Consequently, it has been suggested that terpenes enable plants to send and
receive distinct, targeted, and private information (Rosenkranz et al., 2021).
Adding another layer of complexity, different parts of the plants present distinct emission
patterns and produce different responses to the presence of VOCs. The implications of
organ-specificity are innumerable and can range from maintaining and regulating the
microbiome of flowers (Boachon et al., 2019), to altering root architecture (Ditengou et
al., 2015).

Regarding the role of VOCs, previous research suggests that they are important both at
an individual and a community level. At an individual level, volatile terpenes can
influence plant innate immunity and can alter complex signalling pathways in plant cells.
This allows them to enhance plant fitness, for example via direct interaction with the
TOPLESS transcription factors (Nagashima et al., 2019).

At a community level, VOCs can allow private communication with conspecifics, based
on concentration specificity (i.e. muting signals as a function of distance to avoid
“eavesdroppers”). Generally, these signals have beneficial effects on the fitness of
“listeners”, which has led to competitors adapting to perceive lower concentrations of
terpenes (Rosenkranz et al., 2021). However, volatile cues may also provoke the opposite
effects, by for example turning neighbouring plants’ roots more sensitive against
herbivores (Huang et al., 2019).

Due to their versatility and involvement in countless mechanisms both at an individual

and community level, the importance of VOCs is undeniable. Contrary to popular belief,
VOCs are not futuristic solutions to tackle the looming climate crisis. In fact, they have
been fruitfully used in agriculture for decades, as the base of the push-pull system.

Nonetheless, this landscape is vastly unexplored. To this date, most of the experiments
on this topic have been carried out at laboratories on axenic cultures with no more than
two plants species at once, disregarding the importance of plant communities and other
non-plant organisms. Similarly, microbial interaction with VOCs (production,
transformation, assimilation, and degradation) and other interrupters of this
communication such as pollution require considerable further research.

Currently, the advantages that VOCs can offer in production, are being achieved with
synthetic compounds. The pesticides englobe a huge list of chemical substances such as
fungicides, insecticides or herbicides used for protecting crops (Bernardes et al. 2015).
Its use has spread so much from the modernization of the agriculture -commonly called
green revolution- that in present day some countries are completely dependent on them
for their crop production (FAO, 2020). This represents a global problem because most
pesticides contaminate the water, soil and even the biomass produced (Tudi et al., 2021).
But also represent a problem particularly in the countries than cannot have access to it,
depleting their production capacity and creating more inequality between countries. In
this project we studied the different level of pesticide use in relation to a country's
development to measure the repercussion that our knowledge about VOCs can have not
only in the ecology, but in the country’s development.

The 2030 sustainable agenda adopted by all the United Nations members sets a total of
17 sustainable goals that must be urgently reached (UNDP, 2022). The ecological
profitable agriculture has a relevant role in a huge number of this challenges and the
VOCs can contribute in “greening” the green revolution. This literature collects the
information we have of VOCs till our present days and worldwide statistics with the aim
of showing there is scientific evidence that supports the existence of a potential tool based
in this plant phenomenon to reach the global problems that United Nations consider to be
a priority.

METHODS

The primary source of information was a variety of articles -including those cited-
obtained from PubMed. Nonetheless, TEDx videos (such as Stefano Mancuso’s “Are
plants conscious?”) and resources form the Society for Plant Signalling and Behaviour
were also consulted.

In addition to this, when writing and editing the project, the authors resorted to the
Cambridge and Merriam-Webster online dictionaries.

For the representation of the pesticide’s utilization, the map creator mapchart.net was
used. Some countries were considered non-representational since their medium
temperatures avoid plague proliferation, so they have no need of pesticides (Briere et al,
1999).

Regarding the worldwide data management and statistical analysis, the HDI data was
taken from the United Nations annual report’s tables (UNDP, 2022). The tons of
pesticides used and cultivated hectares were taken from the FAO statistics bank. With
this data, a one-factor ANOVA test was performed with the program EXCEL. Then, the
data was contrasted with a Shapiro-wilk test to evaluate if the distribution of the groups
was normal, and a T-test between each pair of groups to assess whether there were
significant differences or not. Both tests were done using the software RStudio.

FINDINGS AND DISCUSSION

Pesticides in human development and environment.

The United Nations human development 2022 report shows that the human development
index (HDI) is classified as low in the majority of the African countries, medium in the
countries between the tropics, and high or very high in the rest (UNDP, 2022) (Figure 1).
On the other hand, the food and agriculture united nations organization (FAO) have
registered the pesticides use and the cultivated area in each country of the globe. The
results of calculating the tons of pesticides per thousands of hectares are shown on figure
1 and figure 3.

Comparing figures 2 and 3, a correlation

Tons of pesticides per thousands of
hectares vs HDI between use of pesticides and HDI can be
12,00 ab
observed. This hypothesis is corroborated
10,00 ab
by the statistic ANOVA test, which
8,00
6,00 demonstrate there is significant difference
c
4,00 between the use of pesticides in the different
2,00 d groups of countries based on the HDI.
0,00
-2,00 Specifically, the t test indicates that there is

very high high medium low

significant difference between all the groups
Figure 1. Use of pesticides of countries divided by the except for the very high HDI and high HDI
degree of development. Statistically significant
differences are indicated with different letters when p ≤ groups (Figure 1).
0.05.
Remarkably, the less developed countries use the least pesticides because, as figure 3
depicts, these countries are in regions with conditions that favour plague proliferation
(Briere et al., 1999). On the other hand, in the most developed countries there is a social
tendency to use more ecological products than in the developed ones (Heitschmidt et al.,
2004). At the same time, pesticides are more affordable in the most developed countries.
These two circumstances balance out, hence there is no significant difference in pesticide
use between the two most developed groups.
Figure 2. HDI per country. From darker to lighter: very high, high, medium and low. The non-coloured ones correspond
to no data.

Figure 3. Tons of pesticides per thousands of hectares cultivated. Non-coloured countries are those with no data or
non-representational data (Canada, Russia, Sweden, Finland, Iceland and Norway).

This suggest that pesticides play a very relevant role in the inequality between countries,
as those less developed are unable to afford them despite them being essential for their
crops due to their conditions. As a result, they have a lower production ratio, which leads
to these countries being outcompeted in the food market, consequently hindering their
incomes growth (FAO, 2020). So, there is a necessity to improve the production without
the use of pesticides in order to reduce inequality. This is one of the Sustainable
Development Goals and our results about the VOCs elaborated below, show that
advances can be made in this line using them. Another three goals, involving the end of
hunger, of poverty, and the stimulation of economic growth, could also be obtained -
achieving a higher production yield- by relying on VOCs. As they can be directly
impacted by increasing the supply of food, or indirectly by raising the incomes by the
trade with other countries.

Despite of the intensive use of the

pesticides, the reality shows that only
one percent of the total amount
delivered reaches the targeted plants
(Bernardes et al., 2015). The rest is
incorporated into the environment
until its degradation, which,
depending on the environmental
Figure 4. Pesticide behaviour in the natural environment (Tudi conditions and the chemical
et al, 2021).
characteristics, may take years
(Tcaciuc et al., 2018). In the interim, they can only be absorbed by the soil in the area
where they were administered and remain there resulting in severe soil pollution (Fang
Q, 2014). Alternatively, they can move to other environments (Figure 4), leaching
through the soil to the water table or running across the surface to reach wells, rivers, and
seas; they can be transported by gusts of wind; or volatilise, turning into gas that can reach
practically every environment when it precipitates (Tudi et al., 2021). As a result, the
pesticides can be integrated in the environments contaminating it. Numerous studies have
proven the risks for non-targeted organism like microorganisms, birds, aquatic life, the
crops and, indeed, humans (Aktar et al. 2009). To exacerbate this, some of these
compounds, such as synthetic fungicides, reduce the efficacy of the treatment as
organisms like Aspergillus fumigatus develop resistance. Regrettably, the response to this
often is to increase the level of chemicals used. Taking this into account, pesticides go in
the opposite direction of a great number of the SDGs: to guarantee good health and well-
being, to provide good quality water, to develop a responsible production and
consumption style, and to respond to the climate emergency and the loss of biodiversity.

Currently, there is a big pressure coming from the public perception towards pesticides.
Climate change is predicted to affect the frequency and intensity of droughts, which will
trigger a sharp decrease in food production, as well as the progressive aridity of soils.
These conditions will also stimulate the propagation of plant pathogens (Baker et al.,
2000), and therefore, a multiplication of epidemics.

VOCs produced by BCAs.

Biological control agents (BCAs), which are natural antagonists used to mitigate the
detrimental effects of pathogenic (micro)organisms, are decisive to control the pathogen
and/or its detrimental effect. Ideal BCAs should not produce toxic metabolites for both
humans and animals. There are two different ways of action, direct antagonism, which
occurs when the there is a high affinity among the pathogen and the BCA. On the other
hand, indirect antagonism takes place when no physical contact is needed between the
BCA and its target, instead the BCA acts as a “stimulus” to trigger the development of an
unfavourable condition for the microbial growth, leading to a control of the pathogenic
species (Tilocca, 2020).

Among the mechanisms against pathogens that BCAs count with, we can highlight the
production of VOCs. They are produced by all living microorganisms and only some have
a positive effect when controlling phytopathogenic oomycetes, fungi, and bacteria by
means of antimicrobial activity and other cross-walk interactions (Tilocca, 2020).

Push-pull system.

The push-pull system has come to embody a platform for delivery of weed control, plant
nutrition and production of livestock-feed, in addition to pest management for small sub-
Saharan cereal farmers (Picket et al., 2017).

The push-pull system traditionally consists of a “push” plant such as the cattle forage
grass Melinis minutiflora, that is in charge of repelling pests like moths, from the crop of
interest (usually maize). The “push” plant achieves this via the production of the
tetranorterpene DMNT. However, this results in substantially higher parasitism of the
maize by insect larvae, which is where the “pull” plant comes in. The “pull” plant -another
cattle forage grass such as Pennisetum purpureum- releases a high concentration of plant
volatile signals, attracting these larvae. Interestingly, less-genetically-improved species
retain a higher signalling potential, and hence make better intercrops (“support” plants
grown among the plant of interest).
 Furthermore, it has been found that the
signals released by the companion plants
may trigger indirect secondary defence
signalling in certain varieties of maize
plants. When exposed to insect eggs, B.
brizantha altered its’ VOC production,
causing a reduction of herbivory as well
as an increase of volatile attractant
signals to parasitoids (parasites of insect
Figure 5. Attraction of larval parasitoid by maize plant larvae) in the neighbouring maize plants
exposed to signals of Brachiaria plant bearing insect-pest
eggs (Picket et al., 2017) (Figure 5).

In some regions where push-pull is practiced, other intercrops have replaced M.

minutiflora. An example of these are the forage legumes from the Desmodium genus,
which in addition to controlling insect pests, allow the control of parasitic weeds and the
fixing of nitrogen within the system. Recently, drought-tolerant companion crops have
also been employed to accommodate the aridification of cereal-growing regions driven
by climate change.

Consequently, the push-pull system is an excellently advantageous application of VOCs,

which makes the SDGs far more attainable. It is closely related to the no poverty and zero
hunger goals, as it continues to improve the crop yield of smallholder farmers in
impoverished countries, and also provides them with forage for their livestock. Not only
that, but smallholder cereal farmers benefitting from the push-pull system do not normally
buy fertilizers, pesticides, or seed. This goes hand in hand with the responsible
consumption and production goal, as the notorious consequences of fertilizers and
pesticides are concerning at the least. Furthermore, the use of drought-tolerant intercrops
is climate action goal).
Plants’ priming by VOCs.

VOCs participate in the priming of

plants to endure possible exposition
to biotic and abiotic stresses (Figure
6), with a considerable advantage to
other direct protection pathways.
This process does not require highly
energetic metabolic activations (van
Hulten et al, 2006) and provides
plants with memory that allows them
to produce a faster and more robust
answer to future stress. Due to that,
these individuals will be subject to an
improvement in their resistance Figure 6. Possible applications of plant VOCs in agriculture.
Brilli, et al, 2019.
and/or tolerance.

Volatile terpenes can influence plant innate immunity by modulating salicylic acid (SA)
and jasmonic acid (JA) signalling. These phytohormones are recognised as the base of
plant immunity (Tamaoki et al., 2013), as they activate plant defence systems against
different types of pathogen attacks (SA for biotrophic and hemi-biotrophic pathogens, as
opposed to JA for necrotrophic ones) through an unknown mechanism of action.
Monoterpenes have been recorded to induce SA defences (Rosenkranz et al., 2021), thus
carrying out a protective function. On the other hand, the sesquiterpene β-caryophyllene
appears to be strongly correlated with JA immunity. On one study (Frank et al., 2021),
Arabidopsis’ leaves treated with this sesquiterpene induced JA-mediated resistance
against the pathogen P. Syringae, reducing the level of infection.

Isoprenoids, one of the most abundant types of VOCs, are able to produce, by influence
of abiotic stresses, different responses to improve plants answers to these hazards (Figure
6). Among them we can highlight the stabilization of cell membranes, as well as the
quenching of reactive oxygen species (ROS) (Velikova et al., 2011). Moreover, recent
studies (Dani et al, 2016) have suggested that VOCs can collaborate with different
secondary metabolites and hormones synthesised in the pathway involved in the
regulation gradual deterioration of plants. This process, also known as senescence, is
mainly caused by variations in hormones concentrations which cause ROS to accumulate
and harm different cellular components. It is then that VOCs can synergise with the
production of those hormones, inducing an antioxidant response and, because of that,
avoid plants’ aging (Figure 6).

VOCs as possible herbicides.

VOCs also have an important impact in the impediment of competitive plant species’
growth (Figure 6). This competition has been exhibited by different groups, such as
isoprenoids, which include among their effects the absence of seed germination or the
inhibition of root growth. Benefits are obtained by the plants secreting those compounds
as they could act as herbicides by inhibiting weeds from reaching for water or nutrients
and, because of that, increase crop productivity (Puig et al, 2018).

VOCs of Microbial origin.

VOCs of microbial origin in controlling plant pathogens have received a reduced

marginal in the past years, however, it has been highlighted their potential benefits in
agriculture.

Microorganisms have a very close relationship with plants, being able to interact
aboveground, inside them, and in the rhizosphere, underground. VOCs are effective with
just a very small concentration, in order of ng to mg, and being volatile allows them to
diffuse between the soil particles and spread in the atmosphere over large distances from
their emission point. However, it has been demonstrated that the rhizosphere is decisive
in the determination of the specific microbes that reside in it, and therefore, in the types
and concentrations and ratios of VOCs that are secreted. This is mainly due to the
compound that plants secrete through their roots during their nutrition processes.

Studies performed in vitro in Arabidopsis thaliana with VOCs produced by Bacillus

subtilis GB03 (Ryu et al, 2003) showed that they induced not only leaf growth, but they
intensified photosynthesis performance and increased chlorophyll concentrations after
two weeks.

In addition, VOCs are also involved in processes such as the capability to kill plant-
parasitic nematodes, tolerance to drought and salt, and the induction of resistance
mechanisms in plants, preventing them from being colonized by pathogens (Figure 6). A
very well-known example of symbiosis of plants and bacteria are the mutualism of plants
and Rhizobacteria. They are known to produce VOCs blends with antifungal properties
active against pathogenic and non-pathogenic fungi.

Possible challenges to surpass to apply VOCs to agriculture.

There are limitations and drawbacks that should be taken into consideration in order to
advance in the introduction of VOCs in future agricultural procedures.

It is undisclosed if VOCs emissions can be associated with protists, archaea, or other

rhizosphere organisms. Knowing if these organisms have a connection with VOCs would
be crucial. Moreover, once these relationships have been established, effects of these
compounds in the ecosystem should be considered before integrating them in agriculture.

Studies performed regarding this topic have been mainly performed in small scale, Petri
dishes. In these perfectly controlled conditions, VOCs have demonstrated to have an
elevated efficacy in performance. Nevertheless, once they are applied for intensive
agriculture exploitation, more parameters should be considered. Methods regarding micro
and nano-encapsulation of these compounds could emulate their natural segregation to
the atmosphere, but in a more controlled environment which could enhance VOCs’
efficacy. Considering these observations, studies regarding the improvement of VOCs’
target precision should be performed to help achieve a more accurate execution (Brilli et
al., 2019).

VOCs’ high volatility nature, induced by high vapor pressure and low boiling point,
makes them also very unstable in open fields and, therefore, less productive. Moreover,
studies (Blande et al., 2014) have revealed that meteorological circumstances influence
in their performance. Air pollutants are naturally present in the atmosphere, but high
concentrations, mainly induced by human massive industrial production, have dangerous
effects in biotic organisms. Ozone, OH- radicals and NOx have been demonstrated to
negatively impact VOCs’ emission intensity, as these compounds can react with them.
CONLCUSION

VOCs offer great advantages to cope with the heavily demanding agricultural production
and to face the increasing biotic and abiotic stresses. Furthermore, their nature makes
them an eco-friendly alternative to current pesticides that countries could adopt in order
to achieve the SDGs and tackle climate change.

Recently developed technologies (e.g. Proton-transfer-reaction mass spectrometry) could

participate in the phenotyping and quantification of in-vivo emission of VOCs blends.
This data could then be used by breeders to artificially prime plants or to select
phenotypes with the desired characteristics. Moreover, variations in the types and
intensity in which these compounds are emitted could improve crops’ yield. It is also
worth noting that VOCs could offer a competitive alternative to pesticides in areas where
those are not accessible or where their application can cause problems.

However, more time and money investment will be needed to surpass the existing
inconveniences that the introduction of VOCs in extended agriculture presents.
Nevertheless, this could be rapidly achieved as more rigid legislations against current
pesticides are constantly being approved, and therefore, new natural solutions will need
to be studied.

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