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306 8. Medicinal plants from West Africa used as antimalarial agents: an overview
Medicinal Plants as Anti-infectives
PART II
Medicinal plants as
anti-infectives: recent
innovations and regulations
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CHAPTER
9
Mycobacterial quorum quenching and
biofilm inhibition potential of
medicinal plants
Jonathan L. Seaman
1
, Carel B. Oosthuizen
1
, Lydia Gibango
1
and Namrita Lall
1,2,3
1
Department of Plant and Soil Sciences, Faculty of Natural and Agricultural Sciences,
University of Pretoria, Pretoria, South Africa
2
School of Natural Resources, University of
Missouri, Columbia, MO, United States
3
College of Pharmacy, JSS Academy of Higher
Education and Research, Mysuru, Karnataka, India
Introduction
Tuberculosis (TB) caused by Mycobacterium tuberculosis (Mtb) is a complex communica-
ble disease that has saturated the global disease burden and to this day remains the lead-
ing cause of death due to a single infectious organism. Remarkably, TB mortality rates
have exceeded the rate of human immunodeficiency virus and acquired immunodefi-
ciency syndrome, and thus global attention is required to address this pathogen and its
biopsychosocial determinants. Approximately 1.7 billion people worldwide are infected
with TB with an estimated 10 million new cases every year (
WHO, 2019). Although the TB
epidemic is of great concern to world health authorities, 58 million people worldwide
have been cured throughout 200018 and thus the ability of effective diagnosis, treatment
(69 months), and management in curbing the disease burden is entirely feasible. Ending
the TB epidemic has achieved global recognition status and has been included in the
health targets of the Sustainable Development Goals (SDG) as well as the assignment of a
dedicated “End TB” strategy. As with any infectious disease, TB treatment requires a
holistic medical approach that involves a complex interplay between the healthcare envi-
ronment, healthcare practitioner, and the patient. The problem, however, lies in correcting
309
Medicinal Plants as Anti-infectives
DOI:
https://doi.org/10.1016/B978-0-323-90999-0.00008-2 Copyright © 2022 Elsevier Inc. All rights reserved.
the inefficient healthcare practices of the past and striving toward innovative treatment
strategies that are shorter with limited development of antimicrobial resistance. The highly
versatile nature of Mtb, as a pathogen and its ability to be transferred via infectious aero-
sols, coupled with poor healthcare practices such as inaccurate identification and indis-
criminate use of antibiotics, has laid the foundation for an extensive antimicrobial
resistance capacity of Mtb (
Sarathy et al., 2018). This has, in essence, crippled healthcare
infrastructure and has forced scientists and healthcare workers alike to search for novel
strategies to curb this disease. Although 1.7 billion people worldwide are infected with
TB, it is essential to highlight the difference between latent and active infection.
Of the individuals infected with Mtb, only 5%10% will develop active TB (
WHO,
2019
). Latent TB infections should not be ignored as these are a great contributor to bacte-
rial persistence, which has several important implic ations for the development of post-
treatment relapse (
Zhang et al., 2014). Persisting bacterial cells are a population of
dormant bacilli which continue to exist despite the presence of adverse environmental
factors. Latent TB infec tion refers to the presence of infection without the development of
observable disease symptoms. Active TB infection refers to the presence of infection with
the dev elopment of disease symptoms which can be classified as pulmonary (affecting
the lungs) or extrapulmonary TB (disseminated TB) (
Feng et al., 2012). Reactivation of
latent TB is charact erized by the reemergence of disease symptoms in a latently infected
individual. The progression of latent TB to active TB can occur when a latently infected
individual is exposed to the infectious aerosols of an actively infected individual. This
can lead to one of two sc enarios; a progressive active infection or the reactivation of latent
TB, which can be influenced by several risk factors including the immunocompromised
state of the host and/or age (
WHO, 2019). The dynamic nature of TB infections is
represented in
Fig. 9.1.
In order to evade host immune responses and antibiotic bombardment, TB bacilli have devel-
oped a strategy to encapsulate themselves within a protected matrix so that their susceptibility
FIGURE 9.1 Flow diagram of the TB infection
process.
310 9. Mycobacterial quorum quenching and biofilm inhibition potential of medicinal plants
Medicinal Plants as Anti-infectives
and availability to antibiotics and host immune clearance are diminished. These dynamic struc-
tures are often in the form of bacterial biofilms and have been shown to lead to chronic infection
and inflammation (
Erhabor, Erhabor, & McGaw, 2019). Latent TB, and the persistence mechan-
isms, is thus an attractive target for further research as over 60% of infectious diseases treated in
developed countries were due to the presence of biofilms (
Chen & Wen, 2011).
Virulence factor production and biofilm formation are genetically regulated through the
process of quorum sensing, whereby changes in cell density determine the expression of
various pathogenic and nonpathogenic genes (
Adonizio, Kong, & Mathee, 2008). In the
case of Mtb, interference with the quorum sensing cascade, a process known as quorum
quenching, is of beneficial importance as an adjunctive treatment modality for TB biofilm
disruption. Selective pressure for bactericidal antibiotic resistance is minimal when it
comes to quorum quenching strategies as these novel agents do not result in bactericidal
effects, but rather act via the attenuation of virulence (
Kordbacheh, Eftekhar, & Ebrahimi,
2017
). Autoinducer molecules such as N-acyl-homoserine lactone (only Gram-negative bac-
teria), Autoinducer 2, and signaling polypeptides are the chemical signals used as the lan-
guage of communication in cell-to-cell signaling (
Wei et al., 2020). The targets for the
inhibition of signaling include the inhibition of autoinducer synthesis, postsynthetic degra-
dation as well as the disruption of signal reception and subsequent signal transduction
(
Wei et al., 2020).
Although synthetic agents aimed at targeting quorum sensing are currently being
researched and developed, the role of natural products should not be ignored as the pla-
net’s rich biodiversity holds promise for the discovery of a biological gold mine of com-
pounds that can be used to disease. Medicinal plants discovered via ethnobotanical
methodologies are of great interest due to their abundance of bioactive secondary metabo-
lites which function naturally as defense mechanisms; several of which have been shown
to exhibit quorum sensing inhibition properties (
Paul, Gopal, Kumar, & Manikandan,
2018
). This chapter aims to summarize and highlight the potential of medicinal plants and
natural products to act as quorum quenching antimycobacterial agents. In addition, the
current sta te of research has been analyzed to indicate potential gaps within this interest-
ing topic.
Significance of quorum quenching research
There has recently been a transition from The Millennium Development Goals of
200015 to the SDG of 2030. It has been realized that the attention is given to TB and the
effect it has on quality of life has been inadequate and underestimated. This is not a lost
battle, as worldwide attention is producing a myriad of ground-breaking initiatives and
an array of global partnerships that can be used in the fight against the ever-evolving
nature of TB. It is thus a global responsibility to ensure that ground-breaking treatment
strategies are favored in comparison with conventional bactericidal pharmaceuticals as
mechanisms that kill bacteria impose greater selective pressure than those that merely
attenuate their virulence (
Kordbacheh et al., 2017). Among the most notable global TB
initiatives is the “End TB” strategy, as well as the first-ever United Nations (UN) ministe-
rial high order meeting on TB, the outcome of which was a political declaration of
311Significance of quorum quenching research
Medicinal Plants as Anti-infectives
commitment, dedication, and consistency in TB research and novel treatment strategies.
The third SDG illustrates a desire for good health and well-being for all, and the SDG tar-
get 3.3 involves ending the TB epidemic by 2030. Three main areas of interest include
reducing the TB incidence rate, reducing the mortality rate of TB, and finally reducing the
catastrophic losses suffered by individuals and families affected by TB. Reductions of 80%,
90%, and 100%, respectively, are proposed for 2030 when compared to 2015 data.
Formulated in 2014 by the World Health Assembly, the “End TB” strategy is to be used in
conjunction with the SDG and other TB initiatives in the hopes of creating a planet free
from TB and the socioeconomic downfall it entails.
TB has saturated the developing world with eight countries primarily classified as
developing, contributing two-thirds of the world’s global TB burden. These countries
include India, South Africa, Philippines, and Nigeria (
Phetlhu, Bimerew, Marie-Modeste,
Naidoo, & Igumbor, 2018
). There are several socioeconomic and environmental similarities
shared amongst developing countries affected by TB, including the existence of exception-
ally dense urban environments, extensive rural populations, and healthcare that is often
lacking in availability and efficiency. Furthermore, education infrastructure is limited in
these countries, which paves the way for poor compliance to treatment. The disease trian-
gle involves a complex interplay between the host, environment, and pathogen. Each of
these parameters needs to be addressed sufficiently to fully understand the versatile
nature of opportunistic pathogens such as TB.
The significance of a TB adjuvant, focusing on quorum quenching, whose treatment
regime is shorter, whose mechanism of action is selective for biofilm disruption, and
whose availability to the public is improved, could not be more evident.
Current state of quorum quenching research
The raw bibliographic data collected were analyzed using VOSviewer version 1.6.15.
This program allows for a comprehensive and interactive analysis of data acquired from
scientific databases and subsequently generates a schematic representation of various
trends, similarities and differences observed in the research. In the search for novel quo-
rum quenching agents, such analytical interpretations are both relevant and required to
produce high-quality research which is both innovative and justified. An illustration of a
keyword-based search strategy and its subsequent analysis is shown in
Fig. 9.2.
Bibliographic data was collected from a vast collection of journals using a host of scien-
tific databases such as ScienceDirect, Web of Science, as well as Google Scholar. The search
terms used included “mycobacterial quorum sensing,” “mycobacterial quorum quench-
ing,” “mycobacterial biofilm formation,” “mycobacterial virulence factor*,” “medicinal
plant* and quorum quenching,” “medicinal plant* and quorum sensing,” “Mycobacterium
tuberculosis,” “medicinal plant* and biofilm inhibition,” “antimycobacterial medicinal
plant*,” and “phytochemical* and quorum quenching and medicinal plant* and mycobac-
terial quorum quenching.” (* represents singular and/or plural.) The data was acquired
and subsequently analyzed for important trends and research opportunities using a
keyword map.
312 9. Mycobacterial quorum quenching and biofilm inhibition potential of medicinal plants
Medicinal Plants as Anti-infectives
Fig. 9.2 shows that extensive studies have been performed on medicinal plants and their
respective phytoconstituents and that there is a significant link between these studies and
quorum sensing studies. This is advantageous as preexisting research serves as a reference
for information that can be used to optimize protocols and contextualize information. On
the other hand, limitations regarding this observation relate to the test microorganisms
used in these studies. Pseudomonas aeruginosa appears to be the most prevalent microor-
ganism tested in quorum sensing studies along with Chromobacterium violaceum. Although
this is beneficial for studies focusing on P. aeruginosa and C. violaceum, respectively, it lim-
its the extrapolation of data to other species such as Mtb. This may, however, be surpris-
ingly beneficial for novel quorum quenching research on Mtb., as few research provides
greater opportunity for investigation into a previously underexplored topic. One of the
most outstanding observations is the minimal degree of quorum quenching res earch in
general and more specifically on mycobacteria and their associated mechanisms of viru-
lence. As seen in
Fig. 9.2, quorum quenching is a significant distance away from Mtb.,
with no observable link identified by current research initiatives. The fact that such little
experimental evidence exists regarding mycobacterial quorum quenching using medicinal
plants and natural products means that a significant gap in the research has been identi-
fied and that further investigation into this topic will likely yield innovative results which
might be clinically relevant. The virulence of tuberculosis bacilli is associated with their
ability to form biofilms. Although biofilm research has recently been quite concentrated,
the way this mechanism can be inhibited in mycobacteria has drawn little attention.
FIGURE 9.2 Spatial distribution of the current research in quorum sensing and biofilm inhibition.
313Current state of quorum quenching research
Medicinal Plants as Anti-infectives