Sustainability 12 05559 - 2

sustainability
Review
Role of Microorganisms in the Remediation of
Wastewater in Floating Treatment Wetlands:
A Review
Munazzam Jawad Shahid 1 , Ameena A. AL-surhanee 2 , Fayza Kouadri 3 , Shafaqat Ali 1,4, * ,
Neeha Nawaz 1 , Muhammad Afzal 5 , Muhammad Rizwan 1 , Basharat Ali 6 and
Mona H. Soliman 7
1 Department of Environmental Sciences and Engineering, Government College University, Faisalabad 38000,
Pakistan; [email protected] (M.J.S.); [email protected] (N.N.);
[email protected] (M.R.)
2 Biology Department, College of Science, Jouf University, Sakaka 2014, Saudi Arabia; [email protected]
3 Biology Department, Faculty of Science, Taibah University, AL-Madina AL-Munawarah 344, Saudi Arabia;
[email protected]
4 Department of Biological Sciences and Technology, China Medical University, Taichung 40402, Taiwan
5 Soil and Environmental Biotechnology Division, National Institute of Biotechnology and Genetic
Engineering, Faisalabad 38000, Pakistan; [email protected]
6 Department of Agronomy, University of Agriculture, Faisalabad 38040, Pakistan; [email protected]
7 Botany and Microbiology Department, Faculty of Science, Cairo University, Giza 12613, Egypt;
[email protected]
* Correspondence: [email protected]

Received: 6 June 2020; Accepted: 29 June 2020; Published: 10 July 2020
Abstract: This article provides useful information for understanding the specific role of microbes
in the pollutant removal process in floating treatment wetlands (FTWs). The current literature
is collected and organized to provide an insight into the specific role of microbes toward plants
and pollutants. Several aspects are discussed, such as important components of FTWs, common
bacterial species, rhizospheric and endophytes bacteria, and their specific role in the pollutant removal
process. The roots of plants release oxygen and exudates, which act as a substrate for microbial
growth. The bacteria attach themselves to the roots and form biofilms to get nutrients from the plants.
Along the plants, the microbial community also influences the performance of FTWs. The bacterial
community contributes to the removal of nitrogen, phosphorus, toxic metals, hydrocarbon, and
organic compounds. Plant–microbe interaction breaks down complex compounds into simple
nutrients, mobilizes metal ions, and increases the uptake of pollutants by plants. The inoculation of
the roots of plants with acclimatized microbes may improve the phytoremediation potential of FTWs.
The bacteria also encourage plant growth and the bioavailability of toxic pollutants and can alleviate
metal toxicity.
Keywords: floating treatment wetlands; water; plants; microbes; pollutants
1. Introduction
Constructed wetlands (CWs) are purposely designed and constructed systems, based on the
physical, chemical, and biological principles and processes of natural wetlands [1]. The vegetation, soil,
and microorganisms are the main components of a CW that contribute to pollutant removal processes
from wastewater. The associated environmental and economic benefits have established CWs as a
viable option for wastewater treatment [2]. These have been widely applied in the treatment of various
Sustainability 2020, 12, 5559; doi:10.3390/su12145559 www.mdpi.com/journal/sustainability

Sustainability 2020, 12, 5559 2 of 29
types of wastewater, such as municipal, agricultural runoff, storm runoff, and industrial [3–8]. Floating
treatment wetland (FTW) is a novel technology, based on a floating vegetated system, that has unique
abilities to remediate wastewater [9,10]. In FTWs, plants are supported by a buoyant mat or raft that
floats on the surface of the water [11]. The roots of the plants develop below the floating mat, extending
down the water column, and develop an extensive root system beneath the water level [10,12,13].
The development of a widespread and dense root system is necessary for the effective performance of
FTWs [14]. FTWs move freely and thus cover a wider area of water than the emergent root system. In a
FTW system, the rhizomes and dense root structure develop a special hydraulic flow in the water zone
between the mat and the bottom of the water body, and the floating roots act as a filter [15]. This leads
to an effective removal of pollutants from the water due to the availability of the increased surface area
of roots for adsorption and absorption [16]. The roots and rhizomes provide a habitat for microbial
growth and development. The roots and attached biofilms perform different physical and biochemical
processes for the removal of pollutants from the contaminated water [17,18]. In FTWs, pollutants are
removed by three main processes, namely adsorption, sedimentation, and biodegradation [19].
The benefits associated with FTWs have made it a promising ecological remediation technology
in the field of wastewater treatment. These benefits include economic and convenient construction,
no digging/earth moving or extra land acquisition, easy operation and maintenance, floating mats
that are adjustable with a change in the water level, and excellent treatment performance [10,20,21].
Furthermore, the planted vegetation provides economic and ecological benefits such as the use of
vegetation as fodder, providing a habitat for wildlife/aquatic animals, and enhancing the aesthetic
value of the pond [10,22]. Globally, FTWs are being applied to remediate various types of wastewater,
such as eutrophic water, sewage and domestic, storm water runoff, and industrial [23–29].
Microbes have a fundamental role in the remediation of polluted water by FTWs. The bacteria
attached to the roots form biofilms through a repeated proliferation process [30]. The oxygen and
exudates released by the plants create a substrate for microbial growth and colonization on the root
beneath the water level [31]. Thus, along the vegetation, the performance of FTWs also depends upon
the metabolism of the microbial community in water, attached to the roots and floating mats [32–34].
The application of plants in combination with microorganisms in FTWs is an effective and sustainable
approach for the treatment of wastewater [35]. The plant–microbe interaction enhances the efficacy
of FTWs [36]. Although the plant–bacteria interaction plays an essential role in the removal of
contaminants from aquatic ecosystem, the interaction of the plant with bacteria in the FTWs is not well
explored [37].
This paper discusses this important component of FTWs and provides a detailed overview of the
specific role of microorganisms in FTWs. We have summarized the important species of bacteria that
colonize the roots of plants. Furthermore, the specific role of rhizospheric bacteria, endophytes, and
algae in the pollutant removal process in FTWs has been elaborated.
2. Mechanism of FTWs
In FTWs, pollutants are removed from the wastewater by different mechanisms induced by plants,
microbes, and their mutualistic relationships. The presence of a vegetated floating mat in a water body
boosts the pollutant removal efficiency of the system by modifying the physicochemical properties
of the water [38,39]. The physical characteristics of the plant’s roots and the nutrient uptake are
interdependent/interlinked. The type of medium in which the roots exit and the nutrients present
in the medium specify the root’s physical characteristics [9,40]. In general, the roots of plants filter
the particulates present in the water. Nutrients are taken up by the plant’s roots and accumulated in
them, as well as in the parts of the plant above the mat [14]. Most organic pollutants are degraded
by microorganisms present on the roots. However, some of the organic pollutants are taken by the
plants. The organic pollutants can either be accumulated in the biomass of vegetation or degraded by
endophytic bacteria present inside the plants [41,42].
The plants in FTWs contribute to the pollutant removal process by entrapping pollutant particles
in the roots [11,43,44]. The roots of plants act as physical filters, and remove suspended particulate
matter from the water. For an effective removal, there should be dense roots, so that they can act as a
physical filter and a bio-sorbent [15].
The bioactive substances released by the roots have a unique role in the removal of nutrients. These
substances balance pH, and increase the humic content in the water, which results in the adsorption
and/or precipitation of pollutants in the form of insoluble material [15,21]. The neutral pH induced
by the vegetation helps in the settlement of dissolved particulate pollutants [24]. Moreover, these
substances alter the physicochemical condition of water, and increase metal and nutrient removal
and the sorption characteristics of biofilms [45,46]. For example, plants may remove phosphorus by
direct uptake, but the key mechanisms of phosphorus removal are sorption, settlement at the bottom,
and physical entrapment in the roots [47]. The FTWs also inhibit the growth of algal communities by
removing nutrients from the water, thus reducing their population [48].
Roots act as a suitable surface for the formation of biofilms, which enhance the degradation of
organic pollutants and removal of nutrients from wastewater [11]. Root exudates aid in the retention
of microbes on the roots by providing them with nutrients [49]. The roots also provide oxygen to
rhizospheric bacteria for aerobic degradation of organic matter. The biodegradation of organic matter
into simple nutrients occurs when it comes in contact with the biofilm [50,51]. Plants remove these
nutrients through direct uptake [52]. Trapping in the biofilm of the roots of macrophytes is an essential
mechanism for particulate matter removal. Furthermore, roots let microbial colonies assimilate the
carbon compounds and help in the reduction in biological oxygen demand and chemical oxygen
demand [26]. Floating wetlands can work under both aerobic and anaerobic conditions. However,
the nutrient removal under aerobic conditions is higher than under anaerobic conditions [53]. Other
organic compounds are degraded by heterotrophic microorganisms either aerobically or anaerobically,
depending upon the oxygen level in water [54].
3. Important Components of FTWs

FTW is composed of plants that are vegetated in a floating mat. Different types of material are
used as floating
Sustainability mats.
2020, 12, The
x FOR detail
PEER of these important components is described below (Figure 1).4 of 30
REVIEW
Figure 1. Schematic
Figure1. Schematic representation of floating
representation of floating treatment
treatment wetland
wetland and
andpollutant
pollutantremoval
removalprocess.
process.
3.3. Plants
The selection of plant species has a great influence on the pollutant removal process. The
selection of plants depends upon their local availability, the nature of pollutants, and the climate
zone. The plants mostly used to develop FTWs are of Canna, Typha, Phragmites, and Cyperus genera.
3.1. Growth Media

Different types of growth media have been used to provide support to the plants growing on
the floating mat. This growth media can be coconut fiber, peat, soil, bamboo crush, sand, peat rice
straw, and compost [55]. The selection of growth media also influences the pollutant removal process.
For instance, the use of rice straw as growth media improved the total nitrogen removal process by the
formation of thick biofilms, boosting the nitrification/denitrification process [56].
3.2. Buoyancy
In FTWs, different materials have been applied with different natural buoyancies. These floating
materials serve as a platform to fix the plants. The floating mats are made up of different materials
such bamboo sticks, polyester fibers, plastic and foaming sheets [57–59]. The floating material should
be hydrophobic, nutrient absorbent, bacterial adhesive, and with no desorption [15].
Some patent floating mats are also available commercially, such as Beemat®, and Bioheaven® ,
made up of buoyant material with holes for plantation. The wrapped plastic tubes and pipes
manufactured from polyethylene (PE), polypropylene (PP) and polyvinyl chloride (PVC), and PS
(polystyrene) foams are most commonly used for the construction of floating frames and rafts [38].
A natural buoyant material, bamboo, has been found to be a cheap and cost-effective material for the
construction of floating rafts [60].
3.3. Plants
The selection of plant species has a great influence on the pollutant removal process. The selection
of plants depends upon their local availability, the nature of pollutants, and the climate zone. The plants
mostly used to develop FTWs are of Canna, Typha, Phragmites, and Cyperus genera. They have been
widely applied in FTWs for the remediation of different types of wastewater [30,56,61–66]. Some
species of the Poaceae family (Lollium sp., Zizania sp., and Chrysopogon sp.) have been successfully
applied in Italy, China, Singapore, and Thailand to develop FTWs. Some plant species are suitable for
particular regions and have efficiently removed nutrients and other pollutants in a specific climate.
Some other plants such as Phragmites, Carex, Acorus, and Juncus were also successfully applied in FTWs,
and these effectively adapted in several locations. The selection of macrophytes to develop FTWs is
very important for pollutant removal as well as for ecosystem sustainability. The selected plants should
be native, easily available, non-invasive species, perennial, able to thrive in a hydroponic environment
with an extensive root system and aerenchyma [67]. The application of invasive species in FTWs may
result in damage to the ecosystem, and the ultimate cost of habitat restoration may suppress the benefits
gained by pollutant removal. [68]. The characteristics that make these macrophytes ideal for FTWs are
their robust growth tall shoot length, extensive root system, and large aerenchyma in their roots and
rhizomes. Plants with relatively thin fibrous roots have a better performance in total nitrogen removal,
and plants with high total root biomass have a better performance in NH+ -N removal [69]. The root
development depends upon various factors such as species, age, type of plant and concentration of
nutrients, trophic status of water, nature of pollutants, redox conditions, and use of supporting mats
and growth media. A high nutrient load at an earlier plant stage can be harmful to plants and can
damage the root system [70].
Similarly, the high load of toxicants can also hinder the growth of the root by permanently
damaging young plants. The root development of P. australis was constrained up to 40-cm deep after
3 years of plantation due to the toxic effects of digestate liquid fraction. On the other hand, Typha
latifolia and Juncus maritimus did not establish themselves due to the high pollutant load [71].
3.4. Bacterial Biofilm

Bacteria have a unique ability to form biofilms, also known as epiphytic microbes. Biofilm
formation begins with the attachment of free-floating microbes to gas–liquid and solid–liquid interfaces.
Sustainability 2020, 12, x FOR PEER REVIEW 5 of 30
interfaces. These biofilms have a key role in the assimilation of the biogeochemical cycles and the
dynamics of an ecosystem process [72]. In the aquatic ecosystem, aquatic plants are an essential
These biofilms
substrate for thehave a key role in
establishment, the assimilation
growth, of the biogeochemical
and development of biofilms. Aquatic cyclesplants
and the dynamics
release of
oxygen,
an ecosystem process [72]. In the aquatic ecosystem, aquatic plants
essential for aerobic bacteria attached to roots, and stimulate the nitrogen cycle in the roots’are an essential substrate for the
establishment,
surroundings [73,74]. growth, and development
Biofilms are composed of of
biofilms. Aquatic matrix
an extracellular plants release
comprised oxygen, essential for
of polysaccharide
aerobic bacteria
biopolymers, attached
proteins, andto DNA
roots, that
and hold
stimulate thetogether
the cell nitrogen[75]. cycleThein the roots’ surroundings
structural [73,74].
integrity of biofilms
Biofilms are composed of an extracellular matrix comprised of polysaccharide
is obtained by secreted proteins, various types of exopolysaccharides and cell surface adhesions [76]. biopolymers, proteins,
and
The DNA that holdand
development the maintenance
cell together [75]. The biofilms
of these structural integrity
rely on small of biofilms
molecules is obtained by secreted
such as homoserine
proteins, various types of exopolysaccharides and cell surface adhesions
lactones, antibiotics, and secondary metabolites, such as the Staphylococcus aureus matrix, provide [76]. The development and
maintenance
proteins for the of synthesis
these biofilms rely on
of biofilm. small
The moleculesmatrix
extracellular such as homoserine
also lactones,
facilitates the formationantibiotics, and
of adhesive
secondary
protein found metabolites,
anchoredsuch to the the Staphylococcus
as cell wall of S. aureus, aureus matrix,
holding theprovide proteins
cells together for the
within thesynthesis
biofilm byof
biofilm. The extracellular matrix also facilitates the formation of adhesive
interaction with other proteins [77,78]. The extracellular DNA also strengthens the structural integrity protein found anchored
to cell wall For
of the biofilms. of S.example,
aureus, holding
Pseudomonasthe cells together
aeruginosa within
contains the biofilmamount
a significant by interaction
of DNAwith other
to provide
proteins
stability to [77,78].
biofilms The extracellular
[79]. The natureDNA also strengthens
of biofilms and associated the matrices
structuraldepends
integrity of the
upon the biofilms.
types of
For example,
substrates, Pseudomonas
medium, aeruginosa
and growth containsBacillus
conditions. a significant
subtilis,amount of DNA tobacterium,
a Gram-positive provide stability
can make to
biofilms via[79].production
The nature ofof
twobiofilms
differentand associated
polymers: matrices depends
polysaccharide upon the
extracellular types of substances
polymeric substrates,
medium, and growth conditions. Bacillus subtilis, a Gram-positive
and poly-d-glutamate. Both of these polymers contribute to biofilm formation; however, bacterium, can make biofilms
the
via production
contribution of two
of each different
polymer polymers: polysaccharide
is determined extracellular
by strain and prevailing polymeric
conditions [80].substances
The plantsand can
poly-d-glutamate.
also modify the function Both of and thesestructure
polymersofcontribute
the microbialto biofilm formation;
community however,
in their the contribution
rhizosphere [81]. The
of each polymer is determined by strain and prevailing conditions [80].
biodiversity and species of bacteria determine the functions of the biofilms. The biofilm-forming The plants can also modify
the function
bacteria haveand beenstructure
reported of the microbial
as diverse and community
host specific. in their
The rhizosphere
secretion of [81]. The biodiversity
macrophytes and growthand
species
status canof bacteria
determine determine the functions
the bacterial compositionof theof biofilms.
biofilms in The thebiofilm-forming
aquatic ecosystem bacteria
[82]. have been
Moreover,
reported
the bacterialas diverse
communityand host specific. was
of biofilms The found
secretion of macrophytes
to be different thanand growth
those in thestatus can determine
surrounding water
the bacterial
column [37].composition of biofilms in the aquatic ecosystem [82]. Moreover, the bacterial community
of biofilms was found to be different than those in the surrounding water column [37].
4. Microorganisms
4. Microorganisms
Microbial communities have an essential role in the organic and inorganic pollutant removal
Microbial
process and plant communities have an essential
growth promotion in FTWsrole in the
(Figure 2);organic
however, andlittle
inorganic
has been pollutant
explored removal
about
process
specific microbial species in roots and their functions in pollutant removal processes from about
and plant growth promotion in FTWs (Figure 2); however, little has been explored water
specific
[83,84]. microbial
Some bacteria, species in roots
such and their functions
as rhizospheric bacteria,in pollutant
are essentialremoval processes
for vigorous plant from water
growth [83,84].
[85]. The
Some bacteria, such as rhizospheric bacteria, are essential for vigorous
bulk soil is the main source of these microbial populations. However, the rhizospheric bacterial plant growth [85]. The bulk soil
is the main source of these microbial populations. However, the rhizospheric
population is different from the soil bacterial community [86–88]. Similarly, in FTWs, the microbes bacterial population is
different from the soil
can be categorized intobacterial community
biofilm-forming [86–88].and
bacteria Similarly, in FTWs,bacteria.
water column the microbes can be categorized
into biofilm-forming bacteria and water column bacteria.
Figure 2. Role of rhizospheric and endophytic bacteria in plant growth promotion and pollutant
Figure 2.processes.
removal Role of rhizospheric and endophytic bacteria in plant growth promotion and pollutant
removal processes.
In FTWs, the microbial communities mostly originate from ambient water. The amelioration and
scrapping specific to the plants’ roots perform a central part in the formation of specific rhizosphere
microbial communities.
Actinobacteria was found to be a dominant group in the water of FTW systems; however,
Proteobacteria was mainly found in the roots and biofilm samples [89]. In Proteobacteria,
Alphaproteobacteria was found to be abundant in the rhizoplane of plants vegetated in FTWs,
and biofilms were mostly composed of Gammaproteobacteria. The second largest phylum in water
and plant root samples was Cyanobacteria, but it was not found in biofilm samples. In a comparison
of the microbial communities in the roots of Canna and Juncus, it was found that different plants
host different types of microbes in their roots. This difference reveals that plant roots secrete specific
exudates and compounds, which attract specific microbial communities [89]. The plant rhizoplane
in the water column attracts microbes and develops large microbial mass manifests in the shape of a
thick, slimy coat on plant roots.
The presence of autotrophic microbial populations may also depend upon the presence of sunlight,
although, in most cases, the floating mat covers the water surface to minimize the availability of sunlight.
However, some amount of sunlight may be available under the water to support the Cyanobacterial
community. However, the relative abundance of Cyanobacteria in plant root and water samples was
found to be similar. In the roots of FTW plants, the genera of Cyanobacteria (Anabaena and Nostochopsis)
that forms a heterocyst was abundantly observed. This indicates the ability of Cyanobacteria to
associate with the roots of floating macrophytes and survive in available light conditions. In floating
macrophytes, the rhizoplane was found to be enriched with sulfate-reducing bacteria [90]. In FTWs,
even in aerobic conditions, anaerobic zones were found in the rhizoplane of the aquatic plants. These
anaerobic microorganisms belong to sulfate-reducing bacteria and Clostridium. In FTWs, different
sulfur oxidizers and sulfate reducers are essential to make out the sulfur cycle, yield, and depletion of
hydrogen sulfide within the plant rhizoplane [70]. The sulfur-oxidizing bacteria are essential to protect
the plants by the detoxification of reduced sulfides such as hydrogen sulfide.
The FTWs are efficient for nitrogen removal through denitrification by the microbial process.
The nitrifiers are augmented in the aquatic root system of FTWs and responsible for ammonia oxidation.
The Nitrosomonas and Nitrosovibrio (Nitrosospira) were found only on the plant roots of FTWs plants.
The presence of Rhizobium, Bradyrhizobium, Azorhizobium and Azovibrio contributes toward nitrogen
fixation within the FTWs. Several methanotrophs and methylotrophs were also found on plant roots in
the FTWs [91]. These methanotrophs and methylotrophs were also abundant in the rhizosphere of
terrestrial plants, and these were not specific to the aquatic plants. However, these bacteria have a key
role in the rhizoplane of FTWs plants, predominantly under reduced oxygen levels [92].
Proteobacteria were found in the various rhizosphere systems [91,93–95]. The comparison between
FTW plants and terrestrial plants’ rhizosphere microbial communities revealed a distinctive mutualistic
association of aquatic microbes with aquatic plants. Bacillus, a soil bacterial group, was absent in the
rhizoplane of FTWs macrophytes. Similarly, Acidobacteria, the major bacterial group in the terrestrial
plant, was not found in the rhizoplane of an aquatic plant [94,96]. Cyanobacteria were different in the
plant’s rhizosphere compared to the aquatic plant’s rhizoplane [91,93,96].
Pseudomonas has the distinctive capability to degrade several polymers, which are difficult to
demean by any other group of bacteria [97]. Pseudomonas has a dominant role in the degradation
of polyethylene in combination with physical degradation [97]. Pseudomonas was found abundantly
(95.5%) in a sample of floating foam from FTWs. The development of biofilms on floating mats involves
a distinctive mechanism that is different from the formation of biofilm on plant roots and in water
samples [97].
Ammonia oxidizing archaea (AOA) and bacteria can attach to the suspended roots in an autotrophic
water environment [98]. The ammonia-oxidizing archaea and bacteria were found only on the
roots as biofilms. The predominant ammonia oxidizers were ammonia-oxidizing bacteria (AOB)
on the rhizoplane of macrophytes. The Nitrosomonas europaea and Nitrosomonas ureae were well
adapted to NH4 + -N rich environments. However, in the terrestrial ecosystem, Nitrosospira was found
predominantly in AOB communities [98,99].
In a study on three aquatic plants, N. peltatum, M. verticillatum, and T. japonica, the dominant
phylum detected was Proteobacteria, ranging from 37% to 83%, followed by Bacteroidetes (8–38%).
The other phyla found in root biofilms were Chloroflexi, Firmicutes, and Verrucomicrobia at low
frequencies. The dominant bacteria in the phylum Proteobacteria were Alphaproteobacteria, followed
by Betaproteobacteria and Gammaproteobacteria. The other bacteria detected at a low frequency were
Epsilonproteobacteria and Deltaproteobacteria [74].
The class Epsilonproteobacteria was found to be higher in number in vegetated sediment samples
compared to un-vegetated sediments and biofilms [74]. The difference in microbial composition and
epiphytic biomass may be the effect of the difference in plant exudates such as polyphenols and
allopathically active compounds [100]. The plants can increase the quantity and diversity of bacterial
biofilms in the aquatic ecosystem, which ultimately can promote the remediation potential of associated
macrophytes [72].
Epiphytic bacterial communities are diverse and host specific. A similar phenomenon was also
found in other terrestrial and aquatic plants [82,101]. The biofilms attached to roots exhibit particular
niches. The difference in bacterial communities is attributed to the different growth environments such
as the difference in water flow, the availability of light, and nutrients conditions [37]. Additionally,
plant roots, water characteristics, sediment properties, and aquatic animals also influence the nutrient
availability, types, and suitability of the environment for the bacteria. The epiphytic bacteria diversity
and species richness were generally greater on roots than those on stems and leaves. Similarly, the
bacterial species in vegetated sediments were more diverse than in un-vegetated sediments [74].
Similarly, the bacterial population linked with sea grassroots was different from the adjacent bulk
sediment [102]. Thus, the roots of the plant may alter the bacterial community in the surrounding
environment. This difference may be due to the influence of root rhizospheric zones on organic matter
accumulation, chemical exudates, and oxygen concentration [22,103].
Similarly, the biofilm and sediment’s microbial communities were found to be dissimilar from
one another. In biofilms, the percentage of class Alphaproteobacteria was higher than in sediments.
The class Epsilonproteobacteria and Deltaproteobacteria were mostly detected only in sediment.
The parallel findings have been stated by other researchers who investigated the bacterial composition
in the sediments of two lakes in China [104].
4.1. Role of Endophytes

The microorganisms residing in the roots of plants and soil also have a major contribution to
the uptake of metals from the contaminated media. These microorganisms boost the breakdown of
complex organic and inorganic compounds into simple nutrients, mobilize metal ions, and increase
the bioavailability to plants [105–108]. These bacteria, such as rhizobacteria, stimulate the growth of
plants and biomass production, and enhance plants’ uptake of toxic pollutants, and the their ability to
alleviate metal-induced toxicity [109,110]. Endophytic bacteria reside within different tissues of the
plant [111,112], increasing the ability of plants to cope with different biotic and abiotic stresses [113].
Broadly, endophytes perform three major roles in the plant which are its protection from biotic stress,
relieving abiotic stress, and supporting it by providing nutrients such as the increasing availability
of nitrogen, phosphorus, and other essential elements [114]. The prior inoculation of plants with
endophytes can reduce the chances of bacterial, fungal, and viral diseases, and even the damage caused
by insects and nematodes [113,115]. The relationship of endophytes with host plants may be either as
obligate endophytes and or facultative endophytes [112]. In stress conditions, endophytes may help the
plant to relieve stress by the combined action of multiple mechanisms [116]. Direct mechanisms include
siderophore production [117], antimicrobial metabolites [118], phosphate-solubilizing compounds [119],
nitrogen-fixing abilities [120], and phytohormones [42,121,122]. The indirect methods include
bioremediation and biocontrol [123]. It is established that certain endophytic bacteria initiate a system
known as induced systematic resistance in their host. This system is effective against different types of
pathogenic bacteria, by preventing the induced bacteria from causing any visible disease symptoms in
the host plant [113,124]. It is well reported that endophytes stimulate the degradation of xenobiotics and
their supplementary compounds by expressing required catabolic genes. The endophytic bacteria have
evolved various types of mechanisms to nullify the effect of toxic heavy metals and contaminants, such
as the efflux of metal ions, the transformation of pollutants into less toxic forms, and the sequestration
of metal ions on the surface of the cell [125]. Endophytes can also mitigate metal stress by promoting
photosynthesis, anti-oxidative enzyme activities, modifying translocation, and the storage of heavy
metal ions. The inoculation of maize with Gaeumannomyces cylindrosporus significantly improved the
yield and productivity of maize under lead stress [126]. Similarly, Pseudomonas aeruginosa inoculation
increases the cadmium tolerance (Cd) of plants and enhances the accumulation and translocation of
Cd in inoculated plants [127].
The high concentration of toxic pollutants may cause toxicity to macrophytes, thus decreasing
the efficiency of macrophytes to remediate pollutants. The endophytes may overcome this challenge.
Endophytes possess plant growth-promoting (PGP) traits and degradation genes that assists the plant
in handling with several environmental stresses. The endophytes contribute to the decontamination of
mixed contaminants by degradation and heighten the metal translocation by the mutualistic relation of
plants and endophytes [128,129]. A few studies have highlighted the application of endophytes in the
macrophytes of FTWs for the treatment of sewage effluent, textile effluent, polluted river water and
potentially toxic metals [25,130,131]. The major advantage of using endophytes to improve xenobiotic
remediation is that it is easier to genetically modify the microorganisms for maximum pollutant
degradation than the plants. Furthermore, the efficiency of the remediation process can be easily
tracked by the estimation of the abundance and expression of pollutant catabolic genes in soil and plant
tissues. The unique environment of plants facilitates the endophytic bacteria to make large population
sizes due to the minimal competition. The pollutant is degraded by endophyte bacteria in planta, and
eliminates the toxic effect on the plant [113,132].
The application of endophytes in a FTWs system, vegetated with P. australis, improved the
remediation potential of the plant and successfully removed the toxic metals such as iron, nickel,
manganese, lead and chromium from the polluted river water. These inoculated endophytes were
tracked in the root/shoot interior of P. australis, proving their potential role in pollutant removal [131].
The specific strains of endophytic bacteria inoculated to T. domingensis enhanced the remediation of
textile effluent [133]. Similarly, the inoculation of Leptochloa fusca with a consortium of three endophyte
bacteria strains in CWs boosted the efficiency of plants to remediate tannery effluent. This endophytic
inoculation also enhanced the growth of L. fusca, increased the removal of pollutants and decreased the
toxicity of treated wastewater [49].
4.2. Role of Rhizospheric Bacteria

The rhizospheric bacteria in FTWs have a prominent role in the degradation of organic
matter, [134,135], and the translocation of potentially toxic metals [81,136,137]. This bacterial population
differs qualitatively and quantitatively from those found in the bulk soil [138–140]. The microbial
species in soil biota may pathologically infect the roots and rhizosphere biota [141,142]. The plant
roots secrete exudates and metabolites, which chemotactically attract bacteria [143]. The rhizospheric
bacteria of macrophytes in wetlands have a prominent role in the removal of pollutants [144]. The roots
of the plants actually control the microbial colonies in the rhizosphere with the exchange of oxygen,
CO2 , nutrients, and bio-chemicals [145,146]. The iron and ammonia can be oxidized by the oxygen
released from the roots [81,147]. The roots’ microbial populations also have an impact on the emission
of methane, as well as other gases from the wetland system [148,149]. The enzymes and organic acids
released by rhizophytes modify the nutrients and make them available to roots [135].
The roots of wetland plants secrete bioactive chemicals, which favor the development of microbial
communities on roots [150]. The roots can also oxidize and reduce the sulfide present in their
rhizosphere by regulating oxygen concentration, redox potential, and the release of low-nitrogen
exudates such as sugar [151].
5. Role of Bacteria in Pollutant Removal Process
5.1. Nitrogen Fixation

The nitrogen fixation by microbes is a critical natural source of reactive nitrogen in the wetland
ecosystem [152]. The oxygen and organic matter supply from the roots favor the enrichment of
nitrogen-metabolizing microorganisms in the rhizosphere [40,153]. In the rhizosphere of wetland
plants, bacteria transform the nitrogen by ammonification, nitrification, denitrification, uptake, and
the anaerobic oxidation of ammonia by nitrate and nitrogen fixation [154]. The metabolic energy
required for this process is obtained from the oxidation of organic matter and lithotrophy. In wetland
plants, most of the nitrogen metabolism occurs at or near the roots [155,156]. The roots either take up
the produced ammonia or they oxidize it into nitrites and nitrates. That oxidized nitrogen diffuses
to the roots or to denitrifiers, which reduces the nitrate to N2 gas in the absence of oxygen [157].
Microbes perform an N-fixation of non-reactive N2 , and nitrogen is produced [158]. The heterotroph
and autotroph prokaryotes contribute toward the production of a large amount of reactive nitrogen
by nitrogen fixation [152]. The nitrogen fixation by cyanobacteria in wetlands depends upon the
availability of light [152]. The important N-fixing bacterial genera are Enterobacter, Azospirillum,
Pseudomonas, Klebsiella, and Vibrio in wetlands [153,159]. The heterotrophic nitrogen fixer usually
makes mutual symbiosis with the roots and exchanges the sugars from the roots for ammonia that
bacteria produce [152,160]. The nitrogen fixation process took place several times in the planted area of
wetlands relative to the non-planted area, especially in the oxygen-deprived area of wetlands [153,161].
The same bacteria also influence nitrogen fixation and denitrification. Often, these processes take place
concurrently near the roots of macrophytes [162]. The nitrogen-fixing bacteria dwell on the roots or in
the rhizosphere of most of the aquatic macrophytes such as P. australis, J. effusus, J. balticus, Sagittaria
triflolia, Zostera marina [163–165]. Roots also contribute to nitrogen fixation by reducing nitrogen from
their rhizosphere, adjusting the pH level and redox potential [151]. Nitrogen-fixing microorganisms,
such as Azospirillum, reside in the rhizosphere; these stimulate hormones, such as auxins, to influence
the pH and redox potential and boost the nitrogen fixation process [161].
5.2. Degradation of Organic Pollutants

Microbes are known as bio-remediators due to their capability to break down virtually all classes
of organic pollutants [166–168]. Microbes degrade the organic pollutants by a process of co-metabolism.
In this process, microbes in the rhizospheric zone of aquatic and terrestrial plants degrade the complex
carbon-based compounds in order to obtain organic carbon and electron acceptors [169]. In natural
water, the biodegradation rate depends upon the microbial population and amount of xenobiotics [170],
and the numbers of the microbes are heavily influenced by the macrophyte species [171]. Plants give
organic carbon to microbes present in the rhizosphere that assist them to degrade complex organic
compounds [172], such as hydrocarbons and aromatic hydrocarbons [173,174]. Bacteria also release
indole acetic acid (IAA) to improve plant growth [175]. Many bacteria isolated from aquatic plants
also showed pollutant degradation and plant growth-promoting activities [176,177]. The biofilms
attached to aquatic plants are capable of degrading organics such as phenolics, amines, and aliphatic
aldehydes [178]. Additionally, these biofilms are capable of degrading dissolved organic matter such
as polychlorinated biphenyls (PCBs) and atrazine [54,179,180]. The aquatic plant rhizosphere is also
enriched with methanotrophs containing a collection of Proteobacteria, which utilize methane for
obtaining carbon and energy [181]. Methanotrophs can degrade numerous types of harmful organic
complexes [182,183] such as chlorinated ethenes by enzymatic reactions. The Eichhornia crassipes can
remediate eutrophic water by influencing the production of gaseous nitrogen [184,185].
5.3. Removal of Heavy Metals

The rhizospheric and endophytic bacteria have been reported to play a prominent part in the
removal of heavy metals (Table 1). Bacteria promote the removal of metals by their ability to sorb the
metallic ion into their cell walls [186]. Metal uptake by plants can be enhanced by bacteria, which
increase the bioavailability of metals to plants [187,188]. The microorganisms can accumulate heavy
metals with the help of specific metal-binding proteins and peptides such as metallothionein and
phytochelatins [189]. The transcription factors of metal-binding proteins facilitate the hormone and
redox signaling process upon exposure to toxic metals in the context of toxic metal exposure [190].
Cyanobacteria decrease the metal toxicity by the production of proteins that can bind metals [191].
The genetically modified Ralstonia eutropha can reduce the harmful Cd (II) by the production of
metallothionein on the surface of the cell [192]. Likewise, Escherichia coli regulates the accumulated Cd
toxicity by the production of many proteins and peptides [193]. The production of metallo-regulatory
protein is a natural resistant method against arsenic (As) and mercury (Hg) in microorganisms [46].
The metal toxicity affects the performance of the phytoremediation process [194]. Microorganisms
augment and facilitate plants to make heavy metals and antibiotic-resistant proteins [195].
The antibiotic-resistant proteins can reduce the abiotic and biotic stress induced by metals. Some of the
Bacillus sp. strains have the ability to devise a mechanism to alleviate the metal stress by an active
transport efflux pump [194]. The endophytic bacteria also influence the functional and phenotypic
characteristics of the plants in which they reside [196]. Moreover, these bacteria influence the activity
of plant antioxidant enzymes and lipid peroxidation, which support the plant resistance system,
particularly resisting the oxidative stress in the plants caused by heavy metals [197,198]. Methylation
can also be used by a few endophytic bacteria to induce the defense and detoxification of metals. Few
gram-negative bacteria possess the specific mercury-resistant (Mer) operon gene for the degradation of
organic mercurials and reductions in Hg+2 [199].
Table 1. Removal of heavy metals by bacteria.
Bacteria Metal Reference

Lactobacillus delbrueckii and Streptococcus thermophillus Fe, Zn [200]
Acinetobacter sp., Bacillus megaterium and Sphingobacterium sp. Fe, Mn [201]
Anoxybacillus flavithermus Fe, Cu [202]
Leptothrix, Pseudomonas, Hyphomicrobium and Planctomyces Mn [203]
Methylobacterium organophilum Cu, Pb [204]
Herminiimonas arsenicoxydans As [205]
Cd,
Enterobacter cloaceae [206]
Cu, Cr
Pb, Cu,
Acetobacter Mn, [207]
Zn, Co
Cd,
Chryseomonas luteola Co, [208]
Cu, Ni
Ochrobactrum anthropi Cr, Cu [209]
Anabaena spiroides Mn [210]
Ralstonia solanacearum Pb [211]
Proteobacteria and Bacteroidetes Cu [212]
Bacillus cereus Cu [213]
Bacillus licheniformis Pb [214]
Ralstonia solanacearum Pb [211]
Enterobacter aerogenes Cd [215]
Cd,
SPseudomonas azotoformans [216]
Cu, Pb
5.4. Metal Biosorption and Bioaccumulation

Generally, bacteria perform metal ion biosorption into their cell wall by two processes, which are
passive and active [217]. Passive biosorption takes place in the cell walls of living and dead/inactive
bacterial cells, supported by multiple metabolism processes [218]. The reaction between the functional
groups (e.g., amine, amide, carbonyl, hydroxyl, sulfonate, etc.) of the cell wall and metal ions causes
the adsorption of metal ions to the cell surface [106]. In the metal ion binding process, different
mechanisms (e.g., ion exchange, sorption, complexation, chelation and micro-precipitation) may be
involved independently or synergistically [219].
On the other hand, in the active biosorption process, metal ions are up taken by living cells.
The fate of metals that enter the inside of living cells depends upon the organisms and specific elements.
The elements can be bound, stored, precipitated, and sequestered in some specific intracellular
organelles and may be transported to a particular structure [106,220].
The endophytic bacteria exhibited outstanding heavy metal bioaccumulation and detoxification
abilities [59,221]. The plant–bacteria symbiotic relation improves the phytoremediation potential of
plants by the increased uptake of heavy metals due to the secretion of organic acid by bacteria. These
organic acids secrete, by bacterial influence, the pH of the system and increase the bioavailability of the
metal ions to plants [222]. For example, the application of endophytic bacteria, Pseudomonas fluorescens
G10 and Microbacterium sp. G16, on Brassica napus increased the Pb accumulation in plant shoots [223].
Saccharomyces cerevisiae, commonly known as baker’s yeast, is a successful bio-sorbent for the removal
of Zn and Cd due to its ion exchange mechanism [224,225]. Similarly, Cunninghamella elegans has been
proven an efficient sorbent for the remediation of textile effluent enriched with heavy metals [226].
Bacteria also produce biosurfactants and release them as root exudates. These biosurfactants
enhance the bioavailability of metals in the soil and aquatic medium by their interaction and
complexation with insoluble metals [227]. On the other hand, the extracellular polymeric substances,
mainly composed of proteins, polysaccharides, nucleic acid, and lipids, perform a key part in the
complexation of metals and reduce their bioavailability [125]. For example, Azobacter sp. formed
complexes with chromium and cadmium by the formation of extracellular polymeric substances (EPS)
and decreased the uptake of metals by Triticum aestivum [228]. The secretion of different metabolites such
as siderophores and organic acids (including citric acids, oxalic acid, and acetic acid) influences heavy
metals’ bioavailability and their translocation in plants [229,230]. In an earlier study, the inoculation of
the endophytic bacterium (Pseudomonas sp.) improved the plant’s growth and increased the nickel (Ni)
accumulation in the plant [220].
6. Role of Fungi
Fungi perform a potential role in the remediation of heavy metals by increasing their bioavailability
and transformation into less toxic forms [231–233]. Some fungi, such as Klebsiella oxytoca, Allescheriella
sp., Stachybotrys sp., Phlebia sp. Pleurotus pulmonarius and Botryosphaeria rhodina, have the capacity
to bind metals [234]. Fungal species like Aspergillus parasitica and Cephalosporium aphidicola can
remediate lead-contaminated soil by their biosorption process [235,236]. The fungi Hymenoscyphus
ericae, Neocosmospora vasinfecta and Verticillum terrestre showed resistance to Hg and the ability to
transform the toxic state of Hg (II) to a non-toxic form [237]. Fungi of the genera Penicillium, Aspergillus,
and Rhizopus, have proven efficient in heavy metal removal from polluted water [238,239].
Fungi link closely with the roots in wetland plants and have a significant influence on wetland
functioning [240,241]. Root exudates attract fungi toward the rhizosphere. The roots and fungi in
wetland plants make multilevel physical, chemical, hormonal, and genetic interactions, which may
be species specific [242,243]. The rhizospheric fungi community is different than soil communities.
The types and interactions of the fungal community with the rhizosphere may be influenced by plant
species, soil characteristics, climate, type of water, and other microorganisms [244]. The plant–fungi
association in wetland plants performs different key functions such as the emission of metal-chelating
siderophores, denitrification and metal detoxification [245,246]. Bacteria can easily stick to the surface
of the substrate compared to algae due to their smaller size [247]. The other reason for the high ratio of
attachment of epiphytic bacteria to aquatic plants compared to algae is the specific metabolites released
from the plants [184,248].
7. Role of Inoculated Bacteria

It is well established that plant–bacteria synergism is essential to enhance the phytoremediation
potential of plants and ultimately FTWs (Table 2) [49,249,250]. The inoculation of FTWs by immobilized
denitrifiers greatly improved the nitrogen removal from wastewater [61]. Endophytes can be isolated
from and within various plant tissues that include roots, stems, leaves, flower, fruit, and seed [112].
The root is the main source of endophytes, and legume root nodules have a large diversity of
endophytes [251]. Some plants have an underground stem, so, in these plants, stem and root
endophytes may be similar [252]. Bacterial endophytes that were obtained from the shoot of sugarcane
promoted fixation as well as acetylene reduction activities [253]. The inoculation method affects bacterial
colonization, and inoculation should be performed appropriately [254]. Nonetheless, no standard
method is defined for the inoculation of plant roots in FTWs. The two common methods of inoculation
are the inoculation of seeds and the inoculation of soil [252,255,256]. In seed inoculation, the inoculum
is introduced into host plants directly when they are in the seed or seedling stage. The soil inoculation
is done directly in root media or the pot in which the plant is growing. In FTWs, the roots of the plant
are inoculated directly by pouring the inoculum in the water near the root of the plant. For example,
Shahid et al. (2019a) prepared the inoculum of five different rhizospheric and endophytic bacterial
strains and inoculated the roots of plants by directly adding a specific amount of inoculum into
the water [20]. Previously, many attempts have been performed to create an effective partnership
between plant and metal-resistant bacteria in order to effectively treat water contaminated with heavy
metals [250,257,258]. FTWs vegetated with Brachia mutica and inoculated with bacteria were used to
treat sewage effluent and it was found that the concentration of heavy metals, including Cd, Fe, Cu,
Cr, Mn, Co and Pb, decreased significantly from the effluent. The removal of iron was significant
(79 to 85%) [259]. Similarly, in another study, a consortium of hydrocarbon-degrading bacteria was
added into the hydrocarbon-enriched water for its remediation by FTWs [260]. The inoculation of
these rhizospheric and endophytic bacteria was reported to enhance the degradation of hydrocarbons,
and also improved the efficiency of the FTWs.
Table 2. Application of bacteria to enhance phytoremediation potential of floating treatment wetlands.
Bacteria/Bacterial Biofilm Nature of Bacteria Plant Plant–Bacteria Interaction Summary Reference

The removal efficiencies of TN, NH4 + -N,TP, total organic
Ipomoea aquatic and carbon (TOC), Chl-a, total microcystin-LR and extracellular
Bacterial Biofilm __ __ [261]
Corbicula fluminea microcystin-LR were 52.7%, 33.7%, 54.5%, 49.2%, 80.2%, 77.4%
and 68.0%, respectively.
The average removal efficiencies of chemical oxygen demand
Bacteria were mainly attached on the (COD), TN, NH3 -N and TP for Canna indica set-up were 23.1%,
Canna Indica and
Proteobacteria Nitrosomonadaceae fiber filling of floating mat and plant 15.3%, 18.1% and 19.4% higher, respectively, than that of the [262]
Iris pseudacorus
roots setup with only substrate, and 14.2%, 12.8%, 7.9% and 11.9%
higher than Iris pseudacorus. FTWs.
Specific microbial communities were COD was efficiently removed in all systems examined (>90%
Carrying nirS, nirK and
Nitrifying and Denitrifying Unplanted visualized with denaturing gradient removal). Ammonia was efficiently removed by nitrification. [22]
amoA genes
gel electrophores (DGGE) Removal of total dissolved nitrogen was ∼50% by day 28
Carex virgate,
Biofilm performed a key role in the The presence of a planted floating mat with biofilms improved
Cyperus ustulatus,
removal of Cu, P and FSS. Plant roots removal of copper (>six-fold), fine suspended particles
Biofilms __ Juncus edgariae, and [11]
and biofilm interaction enhanced metal (∼threefold reduction in turbidity) and dissolved reactive P
Schoenoplectus
speciation compared to the control.
tabemaemontani
The ammonifying bacteria adhered to
plants roots enhanced oxygen supply
The organic nitrogen decomposition rate was up to 86.50% by
Engineering bacterial to microorganism involved in
Ammonifying bacterial strains Cymbidium faberi adding the strain agent while it was 75.66% without them in the [263]
strain nitrification process and increased
control test group in FTWs
capacity of plants roots to absorb
ammonia nitrogen.
The average removal rates for TN, NH4 + -N, NO3 − -N NO2 − -N,
TP and chlorophyll-a in summer–autumn season were 36.9%,
Adsorptive biofilm Natural Thalia dealbata Combined action of plant and biofilms 44.8%, 25.6%, 53.2%, 43.3% and 64.5%, respectively, effectively [55]
reduced the concentrations of total suspended solids (TSS),
Escherichia coli and heavy metals.
Combined action of plant and
Photosynthetic bacteria __ Vetiveria zizanioids inoculated bacteria improved purifying Efficiently removed TN and TP [264]
effect of FTWs
In the batch reactor, COD was mainly
The removal rate of the COD, NH4 + –N, turbidity, and total
Biofilm Reactor Protozoa and Metazoa Bambusoideae removed by the biofilm on the [265]
bacteria were 11.2–74.3%, 2.2–56.1%, 20–100%
filamentous bamboo
Table 2. Cont.

Phyto-accumulation and
Perchlorate reducing rhizo-degradation were key Pistia showed 63.8 ± 4% (w/v) removal of 5 mg/L level
Acinetobacter sp. Pistia stratiotes [266]
bacterium mechanisms involved in perchlorate perchlorate in 7 days
removal
The average removal rates were 86.32%, 93.60%, 90.12%,
Denitrifying polyphosphate Improved the growth of plant and
__ Festuca arundinacea 72.09%, and 84.29%, respectively, for NH4+ -N, NO3 ¯-N, TN, TP, [267]
accumulating microorganisms biomass
and ortho-P.
The inoculated bacteria showed
Maximum reduction in COD, biological oxygen demand
Acinetobacter, Bacillus cereus and persistence in water as well as
Endophytic bacteria Brachiaria mutica (BOD5 ), TN, and PO4 was achieved by the combined use of [259]
Bacillus licheniformis successfully colonized the root and
plants and bacteria.
shoots of the plants
Analysis showed Ni concentration in leaves were between 23
Metals were found in the root biofilm,
Juncus effuses and 31 µg/g dry matter, and between 113 and 131 µg/g in roots.
Biofilms Natural probably due to microbial respiration [14]
Carex riparia Accumulation of Zn was 45-80 µg/g in leaves and 168–210 µg/g
activity
in roots.
Possessed pollutant-degrading and
Klebsiella sp., Pseudomonas sp. and plant growth-promoting abilities and The average reduction in COD and BOD5 was 87% and 87.5%,
Endophytic Bacteria Typha domingensis [26]
Acinetobacter sp. successful survival of bacteria was and significantly removed heavy metals.
found in plant tissues
Improved nitrification and Significantly higher removal rates of ammonia nitrogen (85.2%),
Nitrifying and
Biofilm Canna indica denitrification process and overall high total phosphorus (82.7%), and orthophosphate (82.5%) were [18]
denitrifying bacteria
removal of total nitrogen observed
The community was mainly
composed of Cyanobacteria,
Proteobacteria, Bacteroidetes, Successfully maintained TN and TP concentration in the river
Periphyton __ Improved its nutrient removal capacity [268]
Planctomycetes, Firmicutes, water at less than 2.0 and 0.02 mg L−1 respectively
Actinobacteria, Chlorobi and
Acidobacteria.
About 89.4% of the TN was removed from autotrophic coupled
Dechloromonas, Thiobacillus and Heterotrophic and Mixotrophic denitrification occurred in
__ floating wetlands, and 88.5% from heterotrophic enhanced [39]
Nitrospira autotrophic auto and heterotrophic bacteria
floating wetlands
Alkane-degrading gene (alkB)
Bacillus subtilis, Klebsiella sp.,
Hydrocarbon degrading Brachiara mutica and abundance confirmed microbial Reduced oil content (97%), COD (93%), and BOD (97%), in
Acinetobacter Junii and [260]
bacteria Phragmites australis growth in plant’s root and shoot and in wastewater
Acinetobacter sp.
water.
The inoculated bacteria showed
successful colonization and survival in Bacterial augmentation enhanced the removal potential
Acinetobacter lwofii, Bacillus cereus, Phenol-degrading
Typha domingensis the rhizosphere, root interior and shoot significantly, i.e., 0.146 g/m2 /day vs. 0.166 g/m2 /day without [269]
and Pseudomonas sp. bacteria
interior of the plant and enhanced bacterial inoculation
plant growth and biomass
Table 2. Cont.

Improved plant biomass and high rate
Plant–bacteria synergism significantly improved the phenol
Acinetobacter lwofii, Bacillus cereus, Phenol degrading of inoculated bacteria survival
Phragmite australis degradation and removal. Highest reduction in COD, BOD, [270]
and Pseudomonas sp. bacteria observed in plant roots, shoot and
and TOC was achieved by bacterial augmentation
water
Achieved successful degradation of
Hexadecane The Inoculated bacteria
Hydrocarbons degradation was recorded up to 92%, COD was
Acinetobacter, Acinetobacter sp., Hydrocarbons displayed highest persistence in the
Leptochloa fusca reduced up to 95%, BOD up to 84%, and TDS up to 47% and [41]
and Bacillus niabensis degrading bacteria roots followed by shoots and then in
alleviated the toxicity
the wastewater and improved plant
growth promoting (PGP) activities
Archaea, anaerobic ammonium High abundance and diversity of The average removal rates of NH4 + -N, NO3 – -N and total
Natural Oenanthe javanica [44]
oxidation (Anammox) bacteria bacteria in planted floating wetland nitrogen were 78.3, 44.4 and 49.7% respectively
Proteobacteria Actinobacteria Bacteria were involved in pollutant Suspended solids, TN, TP, NO3 – -N and COD was 86%, 75%,
__ Eichhornia crassipes [271]
Cyanobacteria, and Rhizorhapis degradation and nutrients removal 80%, 95% and 84%, respectively.
Bacillus subtilis, Klebsiella sp., Persistence of bacteria and expression
Hydrocarbon degrading Typha domingensis and Reduction in hydrocarbon (95%), COD (90%), and BOD content
Acinetobacter Junii, and of the alkB gene in the rhizoplane of [272]
bacteria Leptochloa fusca (93%)
Acinetobacter sp. inoculated plants
Removal efficiency was further
Acinetobacter junii, Pseudomonas Rhizospheric and Phragmites australis and Color, COD and BOD after an 8-day period were 97, 87 and
enhanced by augmentation with [273]
indoloxydans, and Rhodococcus sp. endophytes Typha domingensis 92%, respectively, 87–99% reduction in heavy metals
bacteria and promoted plant growth
Reduced COD, BOD5 , and TOC up to 85.9%, 83.3%, and 86.6%
Consortium of five strains namely
Persistence and survival of inoculated in 96 h, respectively. TN was reduced from 37.5 to 2.07 mg l−1 ,
Aeromonas salmonicida, Bacillus
Rhizospheric and Phragmites australis and bacteria in roots and shoots, and N from 33.3 to 1.23 mg l−1 , and TP from 2.63 to 0.53 mg l−1 .
cerus, Pseudomonas indoloxydans, [20]
endophytes Brachia mutica inoculated bacteria improved the plant Trace metals were also reduced up to 79.5% for iron, 91.4% for
Pseudomonas gessardii, and
growth and biomass production nickel, 91.8% for manganese, 36.14% for lead, and 85.19% for
Rhodococcus sp.
chromium.
Acinetobacter juniistrain, The inoculated bacteria showed
The COD was reduced to 92%, BOD to 91%, color to 86%, and
Rhodococcus sp. strain, and Dye degrading bacteria Phragmites australis persistence in water, roots and shoots [274]
trace metals to approximately 87% in the treated wastewater.
Pseudomonas indoloxydans of inoculated plants of FTWs
Bacillus cerus,
Improved remediation performance of The TN, NO3 −1 and TP contents decreased to 1.77 mg l−1 , 0.80
Cyperus laevigatus,
Rhizospheric and Typha domingensis and inoculated plants, inoculated bacteria mg l−1 and 0.60 mg l−1 , respectively. Additionally, the
Aeromonas salmonicida and [131]
endophytes Leptochloa fusca were found in root and shoots of concentration of iron, nickel, manganese, lead, and chromium
Pseudomonas
inoculated plants in the water lowered to 0.41, 0.16, 0.10, 0.25, and 0.08 mg l−1 ,
gessardii,
Bacteria possessing
These strains were Ochrobactrum
capabilities of Produced biosurfactants and promoted
intermedium, Microbacterium
hydrocarbon Phragmites australis, Typha plant growth. Bacteria showed Reduced COD, BOD, TDS, hydrocarbon content, and heavy
oryzae, Pseudomonas, Acinetobacter
degradation, domingensis, Leptochloa persistent in the rhizoplane, roots and metals by 97.4%, 98.9%, 82.4%, 99.1%, and 80%, respectively, [25]
sp., Klebsiella sp., Acinetobacter sp.,
rhamnolipid production, fusca, and Brachiaria mutica shoots of plants within 18 months.
P. aeruginosa, Bacillus subtilus, and
and plant growth
Acinetobacter junii
promotion.
“__” no data.
8. Conclusions
Microbes, bacteria and algae are the major components of epiphytic microbes, which colonize the
lower surface of floating plants. Bacterial biofilm has a crucial role in the removal of organics, inorganics
and metals in FTW systems. The plant species and pollutant concentration in wastewater influence the
nature and diversity of bacteria. Furthermore, the availability of nutrients influences the metabolism of
bacteria and the pollutant removal efficiency. The rhizosphere and endophytes both have a prominent
role in the pollutant removal process. The rhizospheric bacteria mostly remove the pollutants near
the root system, whereas the endophytes mostly remove the pollutants inside the roots and shoots.
The rhizospheric and endophytic bacterial community also enhances the pollutant removal process by
alleviating the pollutant stress, increasing tolerance towards environmental changes, and regulating
plant growth by direct and indirect mechanisms. The inoculation of plant roots with specific strains of
bacteria also boosts the pollutant removal process.
It is clear from this information that plant–microbe interaction is vital for the pollutant removal
process in FTWs. There is a need to conduct further research to gain a better understanding of specific
microbe and plant interactions and their beneficial role in the pollutant removal process in the aquatic
ecosystem. Environmental factors such as temperature, pH, and the availability of nutrients have
a profound effect on the pollutant removal abilities of microorganisms. These factors need further
investigation to achieve the optimal performance of microorganisms in FTWs. The nature of pollutants
affects the persistence and survival of bacteria and may determine the type of bacterial communities in
a wetland system. Bacteria specific to the removal of particular types of pollutants need to be identified
and isolated for their future application in FTWs. Bacteria that are easy to culture in the lab with
minimal prerequisites, which possess the potential to treat a diverse range of pollutants and can be
augmented with diverse macrophytes in FTWs, need to be widely explored for their use in FTWs.
Author Contributions: The paper was written by M.J.S., S.A., N.N. and M.A. The data were collected and
coordinated by A.A.A., F.K. and M.H.S. The paper was reviewed and revised by M.R., B.A., and M.A. All authors
have read and agreed to the published version of the manuscript.
Funding: The authors are grateful to the Higher Education Commission (HEC) Islamabad, Pakistan, for its support.
Acknowledgments: The authors are grateful to the Higher Education Commission (HEC) Islamabad, Pakistan,
for its support.
Conflicts of Interest: The authors declare no conflict of interest.
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sustainability

Keywords: floating treatment wetlands; water; plants; microbes; pollutants

Sustainability 2020, 12, 5559; doi:10.3390/su12145559 www.mdpi.com/journal/sustainability

3. Important Components of FTWs

3.1. Growth Media

3.4. Bacterial Biofilm

4.1. Role of Endophytes

4.2. Role of Rhizospheric Bacteria

5. Role of Bacteria in Pollutant Removal Process

5.1. Nitrogen Fixation

5.2. Degradation of Organic Pollutants

5.3. Removal of Heavy Metals

Table 1. Removal of heavy metals by bacteria.

Bacteria Metal Reference

5.4. Metal Biosorption and Bioaccumulation

7. Role of Inoculated Bacteria

Table 2. Application of bacteria to enhance phytoremediation potential of floating treatment wetlands.

Bacteria/Bacterial Biofilm Nature of Bacteria Plant Plant–Bacteria Interaction Summary Reference

Bacteria/Bacterial Biofilm Nature of Bacteria Plant Plant–Bacteria Interaction Summary Reference

Bacteria/Bacterial Biofilm Nature of Bacteria Plant Plant–Bacteria Interaction Summary Reference

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