Accepted Manuscript

Aquaponic trends and challenges – A review

Brandon Yep, Youbin Zheng

PII: S0959-6526(19)31381-2

DOI: 10.1016/j.jclepro.2019.04.290

Reference: JCLP 16646

To appear in: Journal of Cleaner Production

Received Date: 27 November 2018

Accepted Date: 22 April 2019

Please cite this article as: Brandon Yep, Youbin Zheng, Aquaponic trends and challenges – A
review, Journal of Cleaner Production (2019), doi: 10.1016/j.jclepro.2019.04.290

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Aquaponic trends and challenges – A review

Brandon Yepa and Youbin Zhengb
a,bSchool of Environmental Sciences, University of Guelph, Guelph, ON N1G 2W1, Canada;
aCorresponding author email address: byep@uoguelph.ca
byzheng@uoguelph.ca
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This article reviews current literature published on aquaponics, a growing technology which uses
aquaculture effluent to grow plants. Aquaponics offers a solution to several sustainability issues,
such as, limited water availability, environmental pollution, increasing fertilizer cost, and
depletion of fertile soils. The commercial and scientific application of aquaponics is growing;
however, there is yet to be a review which holistically analyses scientific literature to indicate
what type of system performs optimally, what will be the most dominant horticultural challenges
as the commercial sector expands, and what direction of aquaponic research will be most
impacting. This review analyzed over 529 publications on aquaponics, from 1978 to 2018.
Through a systematic process, 257 of the most constructive publications were further analyzed
and organized into varying groups based on content. The review found that in the past three 3
years, over 160 scientific articles have been published on aquaponic technology, detailing
numerous trends, technological advancements and challenges associated with the system,
consolidating the expansion of aquaponics and the need for a review. From publications
investigating trends, it was found that decoupled aquaponic systems are becoming increasingly
popular over coupled aquaponic systems, a deep water culture hydroponic component and media
bed component are optimal for commercial and research applications, respectively; Tilapia and
dark leafy vegetables are the most successful species used and Nitrospira may play a more
important role in the aquaponic nitrification process than expected. From publications
investigating challenges, it was found that commercial aquaponics will face difficulty growing
high value flowering crops such as sweet peppers, tomatoes or cucumber, as a result of
suboptimal nutrient ratios in aquaponic solution, specifically the reduced K+, Mg+, and Ca+.
Holistically, it was found that the most important aspect of aquaponics that needs future research
is the role plant promoting microbes play in nutrient uptake. Considering plant growth promoting
microbes are likely the cause of aquaponic plants being able to achieve yields similar to that of
hydroponics, despite nutrient levels being significantly lower, future research in this field can be
paramount to the beneficial use of microbes in all plant production systems.
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Wordcount: 14811

1. Introduction

Aquaponics is the process of growing aquatic organisms and plants symbiotically, in

which the effluent of aquaculture undergoes microbial transformations to be used as a source of

nutrients for plant growth, while nutrient absorption from plants remediates water for

aquaculture. Others have systematically defined aquaponics as ‘…a production system of aquatic

organisms and plants where the majority (> 50%) of nutrients sustaining the optimal plant

growth derives from waste originating from feeding the aquatic organisms’ (Lennard, 2015;

Palm et al., 2018). The term “aquaponics” is a portmanteau of aquaculture and hydroponics.

Aquaculture has been defined as “the farming of aquatic organisms, including fish, molluscs,

crustaceans and aquatic plant” (Food and Agriculture Organization of the United Nations, 1988),

while hydroponics has been defined as “… the production of plants in a soilless medium where

by all of the nutrients supplied to the crop are dissolved in water” (Diver, 2006). Although

hydroponics is considered a well-known technology, the term “aquaponics” is relatively

unfamiliar to the general public, as depicted in a European survey conducted by Miličić et al.

(2017) in which over 50% of individuals admitted they had not heard of aquaponics. Modern day

aquaponic systems have generally taken the form of recirculating aquaculture systems (RAS).

Within these systems, the waste produced by aquatic organisms is filtered through tanks of

naturally occurring microbes, which break down organic compounds and make them available

for plant uptake. The main nutrient conversion occurring is the transformation of ammonia

(NH3+) to nitrate (NO3-), via nitrifying bacteria. The aquaculture effluent, loaded with nutrients,

is then filtered into a hydroponic system where plant roots and microbes are fertilized. The water

is then recycled back to the aquatic rearing tanks, remediated of cumulated nutrients. This

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constructed closed-loop ecosystem has gained significant attention in recent years, as it mitigates

several concerns developing in conventional agriculture.

Primarily, aquaponics has been perceived to have a high water use efficiency, use a

minimal amount of synthetic fertilizer, eliminate the use of pesticides/herbicides or antibiotics,

eliminate the need for soil, simultaneously produce plant and fish, and minimize the release of

aquaculture waste into the environment. Although there is a lack of quantifiable scientific data to

back up some of these merits, the concept of aquaponics alone has allowed it to be regarded as

one of the most sustainable forms of agriculture (König et al., 2018; Palm et al., 2018). For

example, the potential to increase water efficiency in the agriculture industry is significant,

considering that the industry consumes approximately 67% of available freshwater

(Shiklomanov, 1998), with areas such as the Middle East and North Africa having their

agriculture systems consume 90% of the total fresh water available (FAO, 2005). Since most

aquaponic systems are some form of RAS, in which plants remove cumulating nutrients for the

fish, no water is lost, with the exception of fish splashing, system evaporation, plant transpiration

and necessary water removals. RAS alone, use 90 to 99% less water than conventional

aquaculture systems, such as raceways or ponds (Timmons and Ebling, 2010). Aquaponic

systems further improves on RAS’ water use efficiency, as the water typically lost in waste

filtration is utilized by plants. Several studies have found that aquaponic systems typically use

between 0.3-5.0% of total system water per day (Maucieri et al., 2018; Rakocy et al., 2010). In

comparison, some basic recirculating hydroponic systems require complete nutrient replacement

every 2 to 3 weeks (Cooper, 1979; Resh, 1995). Aquaponics also, by definition, utilizes at least

50% of nutrients initially provided through fish feed as plant fertilizer, and therefore supplements

a substantially lower amount of fertilizer than hydroponics and in some cases, supplements no

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fertilizer at all. Reducing fertilizer use in agriculture has a substantial impact when considering

that synthetic nitrogen fertilizer production has been estimated to account for 57% of all

agriculture energy demands (Mudahar and Hignett, 1985) and phosphate reserves are predicted

to be half depleted in the next 60 to 70 years (Oelkers and Valsami-Jones, 2008).

Simultaneously, aquaculture systems only capture 25% of nitrogen in fish tissue, while 75% is

excreted into the environment (Hargreaves, 1998; Krom et al., 1995). Aquaculture operations

require a filter system to remove toxic compounds such as ammonia, nitrite, and suspended

particles from the system. These compounds, if not properly managed, may leach into

neighbouring environments and cause water eutrophication. Considering aquaculture is the

fastest growing agriculture industry in the world and is predicted to provide 54% of the estimated

200 million tons of fish demanded by 2030, this is a sector which can have an immense impact

on the environment (Food and Agriculture Organization of the United Nations, 2018). Finally,

given that aquaponics uses a hydroponic component and therefore does not require soil; its use

can be optimized in controlled environments in urban areas. This can help mitigate production

loss from the land shortage caused by urbanisation.

Aquaponic systems have the potential to avoid some of the major resource inefficiencies

present in conventional agriculture; however, achieving the resource efficiency as mentioned

above, has been largely unquantified in scientific publications. Based on several past reviews,

successful aquaponic operations must consider the impacts of system design (Palm et al., 2018),

system water pH control (Tyson et al., 2011), aeration and filtration technologies (Danaher et al.,

2013), acceptable nutrient ranges (Delaide et al., 2016), pairing of plant and fish species,

microbial populations, nitrogen levels, quantity and type of feed (Endut et al., 2010), pest

management and effective marketing. These factors are the main concern of those managing the

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system. In a broader sense, aquaponics is an even more complex system which involves multiple

disciplines, such as aquaculture, microbiology, ecology, horticulture, agriculture, chemistry and

engineering. As a result, complete and comprehensive reviews on aquaponics are difficult. There

are a few literature reviews already available on aquaponics; however, each of these reviews

generally focuses on a single concept of aquaponics, such as management practices (Tyson et al.,

2011), socio-economic feasibility (Junge et al., 2017), hydroponic systems (Pattillo, 2017),

system design (Palm et al., 2018) and industry trends (König et al., 2018). The most

comprehensive and recent of these reviews was arguably the review by Goddek et al. (2015).

Since this review, there has been over 160 scientific publications on aquaponics. These

publications have cultivated technological advancements, comprehensive industry surveys,

horticulture challenges, and suggestions for future research. Specifically, there has been several

publications focussed on new system designs, evaluation of hydroponic components, successful

plant and fish species, the role of plant growth promoting microbes and the challenges of using

the system from a horticultural perspective. These topics have yet to be holistically summarized

in a literature review. Therefore, this review will summarize current trends in these topics, as

well as provide critical insight on their value to both the commercial and research sectors.

Through the aforementioned topics, this review aims to summarize the following: 1. the

most successful system factors based on current trends in the aquaponic industry, 2. the most

limiting challenges of the aquaponic industry from a horticulture stand point, and 3. the most

impacting research directions for the aquaponic industry.

2. Method

This review was developed using a systematic, strategic and comprehensive literature

review on peer reviewed aquaponic publications. This was achieved by analysing every

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publication retrieved with the keyword ‘aquaponic’ from the oldest publications available (1975)

to the newest (2018) on both the web of science (213 publications, see fig.1.), and the University

of Guelph online library, via Primo (529 publications). Based on the results of this search,

publications with significant value were collected in the reference software Mendeley (255

publications). Publications were considered of significant value if they had the three following

attributes: peer reviewed, contain material relating to current trends and challenges of

aquaponics, and used creditable scientific methods. Publications were then categorized into the

following three categories: trends, horticulture challenges, and advancements. Trends was further

divided into the following sections: system design, hydroponic component, plant species, fish

species, and microflora. Horticultural challenges were divided into nitrogen use efficiency,

nutrients, system water pH and solids. Advancements were later incorporated into either folder

based on an overlap of content. Furthermore, key publications referenced in the literature, which

were not previously collected due to limited terminology of ‘aquaponic’, were additionally

collected in the Mendeley software, expanding the breadth of aquaponic literature reviewed.

From this literature review, it became evident that aquaponic publications have been released at

an exponential rate in the past 3 years (over 160 publications), illustrating the importance of a

literature review on the subject.

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Fig. 1. Web of science citation report for search term ‘aquaponic’.

3. Results and Discussion

3.1 Trends

After conducting the literature review, it became apparent that there were major trends in

the following: types of aquaponic systems, hydroponic components, plant species, fish species,

nitrifying bacteria, microflora and additional species. Each sequential section describes the

specific trends for each category, as well as a discussion on their associated impact on both the

commercial and research sectors.

3.1.1 Aquaponic Systems

With the hydroponic component not taken into consideration (as several different styles

can be used), modern commercial aquaponic systems can be divided into coupled aquaponic

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systems (CAS), and decoupled aquaponic systems (DAS) (Forchino et al., 2017). CAS consist of

one continuous system loop, in which the water has only one direction or outlet in each tank. An

example of a classic CAS would be the UVI (University of the Virgin Islands) aquaponic system

or the Integrated Aqua-Vegeculture System, shown in fig. 2. The majority of modern aquaponic

research articles use varying CAS (Ru et al. 2017; Gullian Klanian et al. 2018). Most scientific

articles are focussed on a certain aspect of aquaponics, as opposed to the system design, and as a

result most CAS used are no more sophisticated than the modern UVI system. In comparison,

DAS utilizes sub-loops within the system in which water can travel in more than one direction in

some tanks, shown in fig. 3. This is done primarily for superior filtration, and increased ability to

manipulate nutrient levels and system water pH (Goddek and Keesman, 2018). The first example

of DAS was the system put forth by Naegel (1977); however, it was not until 2015 that the first

modern day DAS was described by Kloas et al. (2015).

~ Direction of System Water

Aquaculture
Tank

Filter Degassing Hydroponic

Clarifier Tank Bioreactor Tank System

Aquaculture
Tank
Sump Tank
(Pump)

Fig. 2. Example of coupled aquaponic system. Design modified from the University of the Virgin Islands

Commercial Raft System (Rakocy, 2012) and designs presented by (Palm et al., 2018).

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~ Direction of System Water Nutrient and pH

Adjustment Tank
Aquaculture
Tank (Pump)

Bioreactor Two Way

Degassing
Filter Tank
Clarifier
Tank Hydroponic
Aquaculture
System
Tank
Sump Tank
(Pump)

Fig. 3. Example of decoupled aquaponic system. Design modified from decoupled designs presented by

Kloas et al. (2015) and the University of the Virgin Islands Commercial Raft System (Rakocy, 2012).

The first record of a modern DAS was published in 2015 by the Leibniz-Institute of

Freshwater Ecology and Inland Fisheries, Berlin Germany (Kloas et al., 2015). The system was

termed Aquaponic System for emission-free Tomato and Fish Production and was unique in that

it had two separate loops that water could flow between in the system. One loop was a filtration

loop, which allowed water to be mechanically filtered and continuously returned to the fish tanks

without encountering the plants. The second loop (only accessible after going through the first

loop) circulated water between the hydroponic component and a tank where fertilizer could be

added. This allowed water parameters to be changed more drastically for the hydroponic portion

without affecting the aquaculture portion of the system. Plants prefer a hydroponic root zone pH

of 5.8 to 6.2, whereas most aquatic organisms prefer a pH of 6.5 to 9 (Rakocy, 2003; Timmons

and Ebling, 2010). This system allowed pH changes and fertilizer additions to occur without

directly affecting the solution in the fish tanks. After the introduction of this system, DAS began

to gain considerable interest. Schmautz et al. (2016) report modifying their CAS system to a

DAS, while also adding a number of hydroponic components to their system. The advantage to

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using numerous hydroponic components is the increased stability in water quality and the ability

to grow and compare a variety of crops. Goddek and Keesman (2018) reported that there are now

numerous modern DAS in Europe (Tilamur, IGB, and Inagro facilities), some of which have

3500 m2 of grow space (NerBreen, Spain). Although it seems aquaponic systems are moving

towards DAS, at least in Europe, there are some disadvantages to the system in comparison to

CAS. The most predominant challenge to DAS is the increased initial cost to construct. There are

no reports currently available on the economics of DAS versus CAS, understandably so

considering how recent DAS are. However, it is presumed to have higher initial costs, as a result

of increased pumps, pipes and tanks. Furthermore, DAS require more space than CAS, which

takes away valuable space for growing plants. Considering that only 31% of aquaponic facilities

are currently predicted to be profitable and 47% of commercial aquaponics facilities rely on

other agricultural product/service for income, adding more costs to the system would be a

challenge to the commercial sector (Love et al., 2014). A study carried out by Tokunaga et al.

(2015) found that a mid-sized aquaponic farm would have an initial investment cost of $217,078

USD. In this scenario a mid-sized aquaponic farm would consists of a fish tank volume of 76 m3

and a plant growth bed of 1142 m2. The study found that the largest portion of expenses in the

fish system were the cost of tanks and pumps. Engle (2015) similarly found that aquaponic

infrastructure costs were between $285,134 and $1,030,536, for small and large UVI systems

respectively. DAS also require additional fertilizer to be added to the hydroponic loop, as it is

separated to a higher degree from the filtration/mineralization tanks, which supports much of the

nutrient reserves, therefore adding another cost to the system (Goddek and Keesman, 2018). To

overcome some of these disadvantages, novel aquaponic design concepts are beginning to

emerge. Goddek and Keesman (2018) put forth the idea of incorporating a reverse osmosis filter

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(desalination unit) within an aquaponic system to remove salts and concentrate nutrients into

water that can be directly added to a hydroponic loop in DAS. Such a system would allow

cleaner water for the fish, while also concentrating nutrients to be used in a more effective

manner, mitigating the costs of additional fertilizer. However, this system would require a high

quantity of energy to run and would only be economical on a large scale, considering the high

costs of such a filtration unit. Less extreme options to improve filtration methods have also been

suggested. Within both DAS and CAS, modification to the filtration units within these systems

has also been investigated in order to improve management and mitigate solid particles entering

the hydroponic component. Danaher et al. (2013) investigated the use of a swirl separator in

place of a clarifier in attempt to concentrate solids in the clarifier to the bottom and to also

increase the flow of water leaving the clarifier. Their study found no difference in fish and plant

yields or system water quality (pH, DO, etc.) from using the swirl separator in comparison to a

clarifier; however, they found that the swirl separator was simple to install, required minimal

labour and decreased settling time for particulate matter. Future improvements to filters will be

an important consideration, given that increased filtration has been found to increase plant yields

and fish health (Sikawa and Yakupitiyage, 2010). As aquaponics expands into the commercial

sector, entities may quickly adapt DAS over CAS. Entities should be cautioned that it is unclear

if DAS are as sustainable and economical as CAS, based on their higher requirement for space,

infrastructure and additional fertilizer. Entities should investigate both system designs based on

their needs, before implementing either, and future research should be allocated to contrasting

the profitability and sustainability of DAS to CAS, to clarify their advantages and disadvantages

to aquaponic growers. With aquaponics primary selling point of being a sustainable system,

developing commercial systems which are truly sustainable is of utmost value. To add to the

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complexity of aquaponic systems, either CAS or DAS may use varying hydroponic systems,

further affecting the productivity and efficiency of the system.

3.1.2 Hydroponic Components

The three most common hydroponic systems used in aquaponics are varying forms of: i)

deep water culture (DWC) or floating raft technique, ii) media filled grow beds, or iii) nutrient

film technique (NFT) (Goddek et al., 2015; Maucieri et al., 2018). A review on hydroponic

systems in aquaponic publications found that 43% used a media-based system, 33% used DWC,

15% used NFT, and 9% used other less common hydroponic systems (Maucieri et al., 2018).

Less common hydroponic systems include: drip irrigation (Schmautz et al., 2016), ebb and flow

(Knaus and Palm, 2017), and vertical towers/walls (Khandaker and Kotzen, 2018). The review

has found that hydroponic components should be evaluated differently in aquaponic systems than

conventional hydroponic systems. In aquaponics, hydroponic components must handle solution

which is higher in total suspended solids (Monsees et al., 2017b; Rakocy et al., 2004), and is also

dependent on beneficial microbes in enhancing plant nutrient uptake (Bartelme et al., 2018).

Several scientific articles have compared some of the above hydroponic systems against one

another in aquaponic systems; however, their results typically vary from study to study. This

variability is likely due to differences in: aquaponic/filtration systems, environmental conditions

and practical experience in running an aquaponic system.

3.1.2.1 Nutrient Film Technique

A study by Lennard and Leonard (2006) investigating the effectiveness of a NFT system

compared to a media culture and DWC system, found that the NFT system had the lowest yields

of lettuce and consequently removed the least amount of nitrate (20% less efficient at nitrate

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removal). This may be attributed to a lower percent of roots in contact with the aquaponic water,

as opposed to DWC in which all of the roots are completely submerged in the water. The root

mats which occur in NFT systems cause roots to fold on top of each other due to the limited

space provided in the trough (Cooper, 1979). This may limit root contact with the nutrient water.

This conclusion was supported in the recent review by Maucieri et al. (2018), who reviewed 122

articles and found that NFT was the least successful hydroponic component in aquaponic

systems. They found that NFT systems, on average, had lower yields and were less popular in

research articles than media culture and DWC systems. Additionally, NFT has limited surface

area for beneficial microbes, and as a result a biofilter is necessary (Maucieri et al., 2018).

Despite these drawbacks, NFT is still widely used in commercial systems, due to its low initial

costs, simple design and overall ease of operation (Goda et al., 2015; Lennard and Leonard,

2006). Future studies should investigate the effectiveness of NFT against DWC and media grow

beds in larger commercial aquaponic settings as opposed to small scale research systems.

3.1.2.2 Media Culture

Media culture is the use of inert solids (i.e. hydroton, perlite, cocopeat) in grow beds, to

grow plants. Media culture is the most common hydroponic system used in aquaponic research

publications, as it can be used for a variety of plant species and is a feasible option for small

scale research systems (Maucieri et al., 2018; Schmautz et al., 2016). Media beds also provide

more stability for root growth, and as a result larger plants may be better adapted to such systems

(Molovan and Băla, 2015). Another benefit of media based hydroponics, is that the substrates

provide enough surface material for nitrifying bacterial growth, and physical filtration, so that a

biofilter is not needed (Maucieri et al., 2018). Conversely, often times the substrates closest to

the initial inflow of water become clogged with sediment and create an uneven fertigation for the

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rest of the hydroponic grow space or even create anaerobic zones (Mcmurtry et al., 1997;

Rakocy, 2003). This can be overcome by flushing the system with filtered water, or by manually

removing, cleaning and replacing the grow bed medium (Pattillo, 2017); however, this is an

additional maintenance operation which can increase the cost of production. For these reasons,

scaling up systems which use media culture can be difficult to maintain. Media culture should be

recognized as a more suitable hydroponic component for smaller scale aquaponic operations,

while hydroponics components with minimal maintenance needs, such as DWC, should be

recognized for more large scale operations (Palm et al., 2018).

3.1.2.3 Deep Water Culture

DWC, also referred to as deep water technique, or floating raft culture (Pattillo 2017), has

been defined by Vermeulen and Kamstra (2013) as “systems with a water level generally more

than 5 cm deep, with the water kept in circulation, while plants float on the water in Styrofoam

plates or otherwise held in position for the crop production”. The use of floating rafts is the most

common commercial hydroponic component according to an international study by Love et al.

(2015a). DWC is popular amongst commercial growers because of their low maintenance,

maximized root to water contact and ability to support a large number of plants with minimal

materials (Molovan and Băla, 2015; Pattillo, 2017). Lennard and Leonard (2006) found that

DWC removed the most nitrate from an aquaponic system compared to NFT and media culture.

Another advantage to DWC is that in the event of a power outage, plants may survive for up to

two weeks without water flow, whereas a prolonged power outage in NFT or media-culture

would be fatal for plants (Rakocy 2003). In a study by Forchino et al. (2017), it was determined

that DWC had a lower environmental impact than media-culture systems, with the primary

difference being attributed to the inert material required for the grow beds in media culture.

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Although this system requires more water than media culture or NFT, many publications have

recorded improved water use efficiency in DWC systems. Love et al. (2015b) found that a DWC

system used 1% of its total system water per day. No other hydroponic component has been

reported to have such a high water use efficiency, which is significant considering generalized

aquaponics systems are widely promoted for water use efficiencies equivalent to DWC. The

biggest challenge in using DWC is the requirement to aerate (incorporate oxygen) into the water

in the grow beds. Since aquaponic systems depend on a high level of aerobic microbes for

nutrient uptake and plant roots require oxygen to uptake nutrients, there is a very high biological

oxygen demand in the grow beds (Pattillo, 2017). Silva et al. (2018) developed a novel

hydroponic system in which plants were half submerged in DWC while the other half of roots

was exposed to air in between the water and the suspened raft. This system was termed Dynamic

Root Floating Technique. In comparison to standard DWC, their system reduced energy costs by

10.3% in a small scale system, which equated to an 11% cheaper total electrical cost for the full

growth cycle of Brassica rapa sub-species chinensis (pak choi) (32 days). Furthermore, this

study found no difference in dissolved oxygen and total ammonia nitrogen (TAN) in the fish

tanks, or differences in either plant or fish growth parameters, which were the primary concerns

when removing forced air from an aquaponic system. A disadvantage of DWC is the increased

presence of pests. Rakocy et al. (2003) have cautioned that DWC systems may harbour

zooplankton, ostracods, snails and other aquatic pests which may slow plant growth by eating

root hairs and beneficial microbes.

3.1.2.4 Vertical Towers/Walls, Ebb and Flow, Drip Irrigation

The less frequently (9%) reported hydroponic components used in aquaponic systems are

primarily Ebb and Flow or flood and drain tables, drip irrigation and vertical towers/walls.

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Although less common in aquaponic publications, these systems can be effective under certain

circumstances. Schmautz et al. (2016) found that tomato plants under drip irrigation had the

highest yield (18.7 kg/m2) in comparison to NFT (17.5 kg/m2) and DWC (17.4 kg/m2). A downside

to drip irrigation is that it requires a significantly higher amount of energy than DWC or NFT

(Schmautz et al., 2016). Another potential challenge of using drip irrigation with aquaponics is the

buildup of solid particles in the drip lines. Palm et al. (2014) was the only study to report the

successful use of a formal Ebb and Flow in aquaponic publications. Advantages of ebb and flow

include increased aeration with less energy use, increased flexibility in periods of aridity and

saturation, minimal infrastructure and increased surface area for microbial growth (Palm et al.,

2014; Pattillo, 2017). A challenge to these systems would be providing filtered water to the fish

tanks at a constant rate, given the variable rates in flooding the tables. However, the merit to these

advantages and disadvantages remains unclear due to limited published research. The most recent

hydroponic system development for aquaponic systems is vertical towers or walls. The potential

for aquaponics to be a form of urban agriculture has led to the development of systems with a

vertical hydroponic component. At Greenwhich Aquaponics Lab in the UK, Khandaker and

Kotzen (2018) investigated the potential of using a hydroponic tower in an aquaponic system, as

well as potential substrates to be used in the towers. Vertical hydroponics can also be extended to

having multiple layers of NFT, DWC, or drip irrigation as described by Pattillo (2017). Pattillo

(2017) also cautions, however, that vertical systems, specifically towers, are susceptible to

biofouling and clogging.

Based on several articles comparing hydroponic components in aquaponics, the following

conclusions can be made: 1) natural variations, management techniques, and application of

hydroponic components in aquaponic systems make it difficult to derive conclusions on which

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component is optimal; 2) hydroponic components must be evaluated differently for aquaponic

systems to account for higher suspended solids and the dependence of microbes; 3) media beds

are ideal for small scale and research aquaponic systems, based on its ability to grow a variety of

plants and mitigate the use of a biofilter; 4) the DWC component appears to be optimal for

commercial applications, based on its low environmental impact, maximized root to water contact,

consistent record of achieving high yields and ability to match water use efficiencies for which

aquaponics is regarded for.

3.1.3 Fish Species

An international survey conducted by Love et al. (2014) found that out of 257 aquaponic

respondents, 69% used Oreochromis niloticus (tilapia), 43% used ornamental fish and 25% used

Siluriformes (catfish) in their commercial operations. Other commonly reported species of fish

used in commercial aquaponics include: Oncorhynchus mykiss (Rainbow trout), Cyprinus carpio

(Common carp), Lates calcarifer (Barramundi), Micropterus salmoides (Largemouth bass),

Piaractus mesopotamicus (Pacu), Pomoxis (crappies) and Maccullochella peelii (Murray cod)

(Rakocy et al., 2006). Other species of aquatic organisms (i.e. Acipenseridae (Sturgeon),

Salvelinus alpinus (Artic charr)) have been suggested to work well in aquaponic systems but

have not been reported in scientific publications. The primary characteristic for an aquatic

organism to be productive in aquaponics is the ability to tolerate high population densities and

high levels of total suspended solids, nitrogen, phosphorous and potassium (Rakocy et al., 2006;

Timmons and Ebling, 2010). Generally, fish should not be stocked higher than 0.06 kg/L,

although species which can thrive close to this density level are ideal for aquaponics (Rakocy et

al., 2006). The most commonly used, and arguably most successful fish species used in

aquaponics is Nile tilapia, followed by Carp and African Catfish. In the literature review

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conducted here, 43% of published articles used a tilapia species as the primary aquatic organism.

Tilapia thrive in aquaponic systems primarily because they are tolerant to un-optimal water

conditions. El-Sayed (2006) classified tilapia to be fast growing, able to withstand stress and

disease, tolerant to a wide range of environmental conditions, and able to feed on low trophic

levels. Tilapia are naturally low trophic omnivores and are microphagus, meaning they consume

small organisms such as phytoplankton as well as small organic particles (El-Sayed, 2006). As a

result, tilapia can tolerate higher total suspended solids, and nitrite levels up to 44.67 mg/L,

which are normally limiting factors for other aquatic species (Ru et al., 2017). Additionally,

Tilapia do not require a high quantity of growing space on account of their low dissolved oxygen

requirement (can survive at 0.5-1.0 mg/L of dissolved oxygen) and can therefore be stocked at a

higher rate, which is ideal for reaching the nutrient demands of plants in aquaponics (El-Sayed,

2006). Based on aquaponic articles currently available, it is not clear if excretion from different

species have a significant affect on the nutrient levels in aquaponic solution or plant yield. For

example, African catfish, Nile tilapia and Common carp all produced water effluent with nitrate

between 20-42.9 mg/L and phosphorous between 8.2-17 mg/L (Endut et al., 2010; Rakocy et al.,

2004; Roosta, 2014). Concurrently, Knaus and Palm (2017) found that the use of Common carp

effluent resulted in higher yields of cucumber over tilapia effluent, while tilapia effluent resulted

in higher yields of tomatoes. Although it is not clear why tomatoes grew better with tilapia

effluent over carp effluent, it was noted that tilapia had a higher metabolic feeding activity than

carp. This may indicate that tilapia were releasing more feces (nutrients) into the water than carp.

The authors also suggested that using multiple species may be advantageous for developing a

more complete nutrient water profile in the water. The use of multiple aquatic species

(polyculture) in aquaponics to develop improved solution for plant growth has not been

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investigated and carries great research potential, considering the limitation in nutrients for some

plants in aquaponic solution, as is discussed later. In terms of economic contribution, aquaculture

product provides minimal value (Bailey and Ferrarezi, 2017; Engle, 2015), with some aquaponic

studies finding plants provided 3.6 fold the revenue than aquaculture (Rakocy et al., 2004).

Bosma et al. (2017) found that it would be difficult to run a profitable aquaponic operation in the

Philippines with a low value fish such as catfish. They emphasized that the main source of

income would come from the plants, and that only high value fish such as Scortum barcoo (Jade

Perch) would contribute a significant amount to the income of an aquaponic farm; however, high

value fish typically require increased water quality (increased space, lower acceptable suspended

solids) which does not complement most aquaponic systems.

3.1.4 Plant Species

In general, leafy vegetables have been the preferred crop to grow in aquaponic systems,

as they grow well in nitrogen concentrated water, have a short growing period, do not have high

nutrient requirements and there is generally a high demand for them globally (Bailey and

Ferrarezi, 2017). Although flowering crops have a higher economic value than leafy vegetables,

they are more difficult to grow in aquaponic systems due to their heavy nutrient requirements of

phosphorous and potassium, their increased susceptibility to pests and diseases, and their longer

growing cycles (Rakocy, 2003). Bailey and Ferrarezi (2017) found that the UVI system (214 m2

grow space) could make $110,000 a year selling only Ocimum basilicum (basil), whereas they

would only make $6400 selling Abelmoschus esculentus (okra). The same study also indicates

that the value of the crop does not necessarily correlate with profit. In their study they found that

although basil had the highest value per kg ($8.80-11.03 USD), Bibb (Boston) lettuce produced

more income per week per m2 ($7.50-9.20 USD) compared to basil ($3.96-4.96 USD), because

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of increased yield and increased planting density. All fruiting crops (i.e. cantaloupe, zucchini,

and cucumber) did not have weekly incomes per m2 above $1.32 USD. Similarly, most

aquaponic economical studies, which found the system to be profitable, were using leafy

vegetables. Rupasinghe and Kennedy (2010) found an aquaponics farm growing lettuce and

barramundi, had a $22,800 higher annual economic return than the two standalone systems

(Rupasinghe and Kennedy, 2010). The aquaponic farm saved $1,320 on nitrogen and phosphorus

fertilizer, $1,269 on effluent disposal and $3,391 on total variable costs, over the period of a year

(Rupasinghe and Kennedy, 2010). Similarly Adler et al. (2000) estimated that combining a trout

farm producing 22,680 kg per year, with a NFT system growing lettuce and basil, would result in

return of 12.5%, and increased profit through decreased costs of water remediation and increased

revenue from plant production (67% of revenue). This may explain why commercial aquaponic

growers primarily produce leafy greens and herbs in their systems. Love et al. (2015a) found that

commercial aquaponic growers most commonly grew basil (81%), salad greens (76%), non-basil

herbs (73%), Solanum lycopersicum (tomatoes) (68%), Lactuca sativa (head lettuce) (68%),

Brassica oleracea (kale) (56%), Beta vulgaris subspecies cicla (chard) (55%), pak choi (51%),

Capsicum annuum (pepper) (48%), and Cucumis sativus (cucumbers) (45%). Furthermore,

aquaponic systems have expanded to grow plants which can thrive in salt water. An important

plant species that can be grown in salt or brackish water aquaponics is Salicornia persica (Kong

and Zheng, 2014; Turcios and Papenbrock, 2014). Salicornia is a halophyte tolerant of high

salinity and able to absorb high levels of nitrate and phosphate. Furthermore, Salicornia shoots

are a nutritious vegetable high in lipids, omega-3s, and minerals, which are gaining popularity in

the European market (Turcios and Papenbrock, 2014). As aquaponics advanced over the years,

so has the number of successful of plant species; however, minimal scientific articles have cited

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the use of aquaponics to grow flowering plants. Future studies should investigate the

effectiveness of aquaponics for the floriculture industry.

3.1.5 Nitrifying Bacteria

Arguably one of the most important organisms in any aquaponic system is the nitrifying

bacteria. Nitrifying bacteria primarily converts TAN into nitrate (NO3-), a form of nitrogen plants

can readily uptake (Canfield et al., 2010). This nitrogen conversion occurs in a two-step bacterial

process. Before the bacterial process, TAN must first be made available in the water. TAN may

be excreted by fish in the form of urine (urea) and feces, of which nitrogen makes up between

10-40% of, or released through gills as ammonia (Wongkiew et al., 2017a). Once in the water,

TAN can be utilized as an energy source for ammonia oxidizing bacteria, while NH4+ itself can

be taken up by plants. Ammonia oxidizing bacteria produce nitrite, which is simultaneously used

as an energy source for nitrite oxidizing bacteria, where it is converted into nitrate. Nitrifying

bacteria may grow throughout the system, but are typically more concentrated in biofilters, if

present. Nitrification is optimal when the temperature is between 25-30 C, the pH is between 7

and 9 (optimally 7.8 according to Antoniou et al., (1990)), and oxygen is below 20 mg/L

(Rakocy et al., 2006). Nitrite at high levels (0.25-1 mg/L) (Jiang et al., 2014) can enter the blood

stream of aquatic organisms and oxidize the iron in hemoglobin molecules, changing it from the

ferrous state to the ferric state (Timmons and Ebling, 2010). This creates methemoglobin, a toxic

compound which can turn the blood brown and result in the lethal Brown-Blood Disease. Nitrate,

however, is a generally non-toxic compound which can be found at levels exceeding 1000 mg/L

in freshwater environments without negative effects on aquatic life (Timmons and Ebling, 2010).

In aquaponic systems, nitrate has been reported to be harmless at concentrations of 150-300

mg/L (Graber and Junge, 2009; Hu et al., 2015). Goddek et al. (2015) summarized that based on

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previous studies, the three major nitrifying bacteria in aquaponic systems are Nitrobacter,

Nitrosomonas, and Nitrospira. However, the notion that Nitrosomonas is the primary nitrite

oxidizing bacteria and Nitrobacter is the primary ammonia oxidizing bacteria, is quickly

changing. A study released by Schmautz et al. (2017) investigating microbial communities in

varying locations of an aquaponic system, found that Nitrospira made up 3.9% of the microbial

community in the biofilter, while Nitrobacter and Nitrosomonadales only made up 0.11% and

0.64% respectively. Similarly, Bartelme et al. (2017) found that in the biofilter of a RAS,

Nitrospira amoA was the most populated nitrifying bacteria, while Nitrobacter populations were

not detected. They also found that Nitrospira amoA populations remained stable over time

despite competing species (ammonia-oxidizing archea), indicating a stable co-existence between

microbes. This was a significant discovery considering that Nitrospira is believed to include

some strains which can convert ammonia to nitrate in a one-step process (Daims et al., 2015). It

is known that complete nitrification (i.e., NH3+ to NO3-) in a one step process, is an energetically

more favourable reaction than the two nitrification steps carried out by ammonia oxidizing

bacteria and nitrite oxidizing bacteria. However, previous studies have found the phenomenon of

nitrification only occurring in the less favorable two step-process. This mindset has changed

recently, as Daims et al. (2015) has found that a strain of Nitrospira, Candidatus Nitrospira

inopinata, is a ‘comamox’ (complete ammonia oxidizer). Furthermore, they found this strain able

to thrive in the following areas: biofilms, flocs and microcolonies of low substrate concentrations

(Daims et al. 2015). It has also been found that Nitrospira are more prevelant over Nitrobacter in

environments with lower nitrite and ammonium concentrations, due to a lower half-saturation

constant (Ks) (Blackburne et al., 2007). This may indicate that Nitrospira will have higher

populations in aquaponic systems which have an efficient nitrifcation process, as such

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environments should have low nitrite levels and high nitrate levels. Future studies should

investigate the varying strains of Nitrospira in aquaponic biofilters, and determine the effects of

nitrogen use effiency when such strains are present. Furthermore, within these biofilters,

nitrifying bacteria often form biofilms or relationships with heterotrophic organisms, such as

protozoa, micrometazoa and heterotrophic bacteria (Timmons and Ebling, 2010). These

organisms may play a role in enhancing nutrient availability to plants.

3.1.6 Microflora

Bartelme et al. (2018) outlined that plant growth promoting microorganisms (PGPM) or

microflora may play a major role in the plant’s ability to uptake nutrients in an aquaponic

system. Although many studies have investigated PGPM in soil environments, there are minimal

studies published on PGPM in soilless environments, due to the sterility and lack of necessity for

PGPM in hydroponics. Therefore, there is an immense opportunity to investigate PGPM in

aquaponics. Bartelme et al. (2018) summarized that species such as Pseudomonas, Bacillus,

Enterobacter, Streptomyces, Gliocladium, or Trichoderma could increase nutrient availability for

plants. For example, the addition of Pseudomonas fluorescens Pf-5 is known to increase

siderophore production in roots found in soil. Siderophores are structures which bind to iron and

facilitate its transport into plant roots. Therefore, Pf-5 may be a valuable PGPM in remedying

common iron deficiencies in aquaponic systems (Bartelme et al., 2018; Goddek et al., 2015). It is

also possible that arbuscular mycorrhizae fungi play a key role in phosphorous absorption in

aquaponic systems. Arbuscular mycorrhizae fungi has been shown to increase phosphorous

uptake in low phosphorous environments, which is significant considering aquaponic systems

can often have low concentrations of phosphorous in the water (Akiyama et al., 2005; Brunno da

Silva Cerozi and Fitzsimmons, 2017). Schmautz et al. (2017) sampled different locations of an

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aquaponic system and found that although similar microorganisms were present in all locations,

their populations differed. It was found that the biofilter had high levels of Rhizobiales and

Actinobacteria, while roots had high levels of Burkholderiales, Flavobacteriales, and

Pseudomonadales. Pseudomonas spp. is a significant microorganism because it is capable of

producing antimicrobial properties as a method of expansion, which simultaneously protects the

surface area they are growing on from disease, such as root rot caused by Pythium (Avis et al.,

2008; Schmautz et al., 2017). This may be a factor in the ability of aquaponic plants to mitigate

waterborne diseases. It has been suggested that PGPM of different species can have a synergetic

effect on plant growth when more than one are present at the same time; however, future studies

are needed to clarify these mechanisms (Avis et al., 2008). Speculation of increased plant

performance from PGPM has led to the development of PGPM culturing. Cerozi and

Fitzsimmons (2016) predicted that plant performance could increase with increased bacillus

strains, as bacillus is known to make phosphorous more available through mineralization and

solubilisation of precipitates. They found that adding “Sanolife® PRO-W; 5.0 × 1010 CFU g−1,

INVE”, twice a week to their aquaponic system at 0.02 g of product per liter of water,

significantly increased plant yields and phosphorous content of lettuce, when compared to lettuce

grown in an aquaponic system without added bacillus. Additionally, it was found that in the

system with bacillus inoculation, ammonia decreased and nitrates were higher, signifying the

utilization of nitrogen and possible promotion of nitrifying bacteria (Cerozi and Fitzsimmons,

2016). Similarly, Zou et al. (2016b) added nitrifying bacteria to improve the nitrogen use

efficiency (NUE) of their aquaponic system. They found that adding B103 (BIOZYM, USA)

(composed of nitrobacteria, denitrifying bacteria, bacillus, lactobacillus, and actinomycetes) on a

weekly basis resulted in lettuce yields improving by 15% and NUE increasing by 4.4%.

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Considering multiple studies on plant improvement through microflora addition, it is suggested

that microflora play a key role in the success of plants in aquaponic systems. There is however,

few studies which investigate the key mechanisms and species of microflora, which allow this to

happen in aquaponic systems. Future research on such matters could have significant value in

terms of application for increased crop production with lower requirements of nutrients.

3.1.7 Additional Species

Additional aquatic organisms may also be added to different parts of the system for

further benefit. Rakocy et al. (2006) reported including fingerling tilapia in the clarifier and

associated pipes to remove solid buildups which may lead to clogging. Fang et al. (2017)

attempted to improve NUE by incorporating microalgae bacteria (consortia) in the bioreactor of

an aquaponic system. The algae aquaponics system had an increase in NUE by 13.8%, an

increase in dissolved oxygen in the bioreactor, and lower N2O emissions. Microalgae provides

O2 for bacteria to mineralize organic nutrients, while simultaneously the CO2 produced by

bacteria supports microalgae photosynthesis. Microalgae has also been shown to reduce

ammonia levels in the water compared to bio-filters containing nitrifying bacteria (Gilles et al.,

2014). This may be a more ideal option for those focussing on the aquaculture portion of the

system for income, as microalgae can be used as a fish feed or as a biofuel, while simultaneously

removing high levels of nitrogen from the system without taking up a high quantity of space.

Another method of increasing the NUE of aquaponic systems is adding new species to mitigate

dissolved inorganic nitrogen. Marques et al. (2017) reported using polychaetes (worms) in a sand

filter to decrease organic solids and unavailable nitrogen. They found that with the addition of

Hediste diversicolor (Ragworm) infused in the sand filter, organic material was decreased by

70% and dissolved inorganic nitrogen was decreased by 67%.

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3.2 Horticulture Challenges

Even in well designed and managed aquaponic systems, there are some fundamental

challenges in using the system for plant production. After the literature review it became evident

that there are four main areas of aquaponics that have been suggested as limitations to optimizing

plant production and resource use efficiency. They are as follows: nitrogen use efficiency

(NUE), nutrient limitations, pH limitations and solid accumulation in the system water. The

specific challenges these categories pose, are described sequentially, followed by a discussion on

their role in future commercial and research based applications in horticulture.

3.2.1 Nitrogen Use Efficiency

As a result of a highly promoted nitrification process, nitrogen (NO3-, NH4+) available for

plant uptake is abundant in most aquaponic systems; however, the efficiency at which nitrogen is

assimilated has not been well-recorded in earlier aquaponic publications. Speculation over the

true NUE of aquaponic systems began to arise after the system became regarded as highly

resource efficient. A number of studies such as Zou et al. (2016), Fang et al. (2017), and

Wongkiew et al. (2017a, 2017b), evaluated the NUE of aquaponic systems, as well as methods

that could be used to increase their efficiency. Studies found that on average, aquaponics has a

NUE between 34.4 and 56.6% (Fang et al., 2017; Zou et al., 2016b). In comparison, 50% of

global fertilizer is applied to rice, wheat and corn, which typically have a NUE below 40%

(Canfield et al., 2010). Other studies have estimated that conventional agriculture has, on

average, a NUE of 50% (Eickhout et al., 2006). This would indicate that aquaponics has a similar

or possibly lower NUE than conventional agriculture, despite being renown as a superior

resource efficient system. Fish assimilate 20-30% of nitrogen provided in fish feed on average,

indicating that 70-80% of nitrogen is released into the water (Hargreaves, 1998; Krom et al.,

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1995). Of the 70-80% of nitrogen released in the water, only 10-37% is typically assimilated by

plants (Wongkiew et al., 2017b; Zou et al., 2016b). The remaining 43.4-65.6% is lost through

nitrous oxide (N2O) emissions (1.5-1.9%) (via denitrification, nitrification, and anaerobic

ammonium oxidation -‘anammox’), ammonia volatilization, water accumulation, sediment

accumulation, solids removal, and assimilation from other microorganisms (Hu et al., 2015; Zou

et al., 2016b). Although nitrogen deficiency is not a typical problem in aquaponic systems,

increasing the NUE appears to be an area of interest, based on literature reviewed. NUE can be

improved by increasing the frequency of feeding (increasing feeding frequency from 8 to 12

times resulted in a 4.9% increase in fish growth and an 11% increase in plant growth) (Liang and

Chien, 2013), decreasing daily feed input (decreasing feed input from 50g/plant to 35g/plant

increased NUE by 18.8% in a lettuce system, and by 22.1% in a pak choi system) (Wongkiew et

al., 2017b), using plant species with higher root biomass (i.e. tomatoes have a NUE 6.9% higher

than pak choi) (Hu et al., 2015), adding nitrifying bacteria (Zou et al., 2016b), using media beds

(Zou et al., 2016b), and using a lower pH (6.0) (Zou et al., 2016a).

Although current aquaponics do not appear to have a NUE superior to that of other forms

of agriculture, the location and form of unused nitrogen is different from that of conventional

agriculture. For example, nitrogen loss from N2O represents one of the more detrimental losses

of nitrogen in any agriculture system. Although minimizing N2O has been of interest in

aquaponic research, its production in aquaponic systems is miniscule in comparison to other

contributors. In the US, in 2016, N2O emissions were equivalent to approximately 390 million

metric tons of carbon dioxide equivalent (EPA, 2010). 77% of these emissions originated from

agriculture soil management, where N2O was released through poor/excessive fertilizer

application and burning of agriculture residues (Eickhout et al., 2006; EPA, 2010). Aquaponics,

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in comparison, does not typically use nitrogen fertilizer, and nitrogen cannot be lost from

weather events. Most of the un-assimilated nitrogen in aquaponics is retained in the water,

sediment or microorganisms. Furthermore, if the most N2O emitting aquaponic system published

by Zou et al. (2016b) (emits 508.2 ugN/m2/h ~ 44,520 gN/ha/year) was scaled up across every

hectare of arable land in the United States (152,262,500 ha), the N2O emissions would be equal

to 6.7 million metric tons N/year, significantly lower than the current 300 million metric tons

produced by current agriculture (EPA, 2010). In appears that the need to improve NUE in

aquaponic systems is based on the system not having a superior NUE to other forms of

agriculture, despite being regarded as a highly efficient system. However, the release of nitrogen

compounds into the environment is still minimal in comparison to conventional agriculture. In

summary, aquaponics NUE can be improved; however, NUE does not appear to be a limiting

challenge for the expansion of commercial aquaponics. A more crucial limiting challenge in

aquaponics is the ratio of nutrients available for plants to uptake.

3.2.2 Nutrient Limitations

In aquaponic systems the four most common limiting essential nutrients for plants in the

solution are Ca, K, Mg and Fe (Rakocy, 2003; Villarroel et al., 2011). Since fish have minimal

requirements of many metal ions (Fe, Mn, Mg, Cu), and lower requirements for K (only 1% of

composition) (Seawright et al., 1998), these nutrients are accordingly low in fish feed, and

therefore low in aquaculture effluent (IAFFD, 2018; Rakocy, 2003). The major ingredient in

most fish feed is fish meal (Brunnoda da Silva Cerozi and Fitzsimmons, 2017). Fish meal is full

of amino acids (organic forms of nitrogen) and phosphorous, but lacks K and several

micronutrients (Fe, Mn, Cu) required by plants (IAFFD, 2018; Savidov et al., 2007). These

nutrients are made further unavailable for plant uptake, through chemical antagonisms in the

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system and non-optimal system water pH. To prevent deficiencies in these nutrients, some

aquaponic systems supplement synthetic salts (i.e. potassium hydroxide) into the system water

(Rakocy, 2003), or apply a foliar spray (Roosta, 2014). Although aquaponic systems aim to rely

as little as possible on synthetic fertilizer for plant growth, it is possible to make up nutrient

deficiencies and increase yield significantly with minimal synthetic fertilizer application.

Of the limiting nutrients in aquaponic systems, K has been reported to be one of the more

significant. Graber and Junge (2009) found that aquaponic water had on average 45-fold less K

than hydroponic solutions. Considering that K is the fifth most up taken nutrient, represents as

high as 10% of plant tissue (Seawright et al., 1998), and plays an essential role in nutrient

transport (specifically Fe and NH4+ transport), enzyme activation (phosphoenolpyruvate

carboxylase (PEP-case) glutamine synthetase (GS)), and moderating osmotic potential,

potassium deficiency can be detrimental to plant growth and yield (Gajdanowicz et al., 2011;

Gierth and Mäser, 2007). Two methods of overcoming K deficiencies are incorporating K into

the system water as a salt, or applying to plants as a foliar spray. Roosta (2014) found that a K

foliar spray on parsley increased yields by up to 60.7% as compared to parsley without a foliar

spray. Gullian Klanian et al. (2018) found that K added to the solution improved tomato fruit

yield, while K in the foliar fertilization improved plant growth. Alternatively, UVI systems

increase K in the water by adding potassium hydroxide (KOH) (Rakocy et al. 2004b, 2006).

KOH has the secondary purpose of increasing or maintaining a neutral pH in acidic aquaponic

systems. K also affects the uptake of other nutrients, specifically Fe. Fe is a nutrient already

typically deficient in aquaponic systems; therefore, its supplementation is crucial if K is also low.

Hydroponic crops typically receive Fe in an available form such as soluble Fe-EDTA or Fe-

EDDHA at a concentration between 2 to 5 mg/L (Goddek et al., 2015; Radzki et al., 2013). In

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aquaponics Fe can be found in the water at concentrations of 0.2 to 2.5 mg/L (Bartelme et al.,

2018; Goddek et al., 2015). In addition to the minimal input of Fe into aquaponic systems, Fe

can be made further unavailable through the speciation of ferrous iron (Fe2+) (highly soluble

across rhizoplanes of roots) to ferric iron (Fe3+) (less soluble across rhizoplane of roots), via

ionic reactions and hydroxyl radicals in the system water (Bartelme et al., 2018; Rose and Waite,

2002). Roosta and Mohsenian (2012) investigated the impact of varying Fe sources as a foliar

spray for peppers in an aquaponic system. They found that a foliar spray containing FeSO4 led to

higher vegetative and reproductive growth in peppers, compared to foliar spays containing Fe-

EDDHA, and Fe-EDTA. Alternatively, the UVI systems add chelated Fe2+ into their system

water at a rate of 2 mg/L, every three weeks for their 214 m2 growing area (Rakocy et al., 2010).

Rakocy et al. (2003) suggests adding iron as Fe-DTPA as it remains stable at a pH of 7.0.

Additionally, the fish species present in the system will dictate whether Fe addition is suitable.

For example, O. mossambicus fingerlings have a LC50 of 8.00–9.38 mg/L of Fe, whereas C.

carpio has a LC50 of 1.22–2.25 mg/L of Fe for 24-96 hrs (Mashifane and Moyo, 2014). Similar

to Fe, Mn, which may also be deficient in aquaponic systems, can cause chlorosis when deficient

(Buhmann et al., 2015). Buhmann et al. (2015) found that Mn addition only increased plant

growth when it was included with Fe-EDHHA in a saltwater aquaponic system. Another nutrient

commonly supplemented in aquaponics is Mg. Mg has been reported to be concentrated around 4

mg/L in aquaponics (Villarroel et al., 2011), which is significantly lower than 24-50 mg/L found

in lettuce hydroponics (Sonneveld and Voogt 2009; Resh 2012). Mg can be supplemented as

dolomite (CaMg(CO3)2) which also doubles as a base to raise the pH in acidic systems (Rakocy,

2003). There are limited studies on the severity of Ca, Mg or Mn deficiencies on aquaponic

plants. Future studies should investigate if these nutrients are causing growth limitations, and if

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so, the effects of additional Ca, Mg or Mn in the form of a foliar spray or system addition should

also be studied.

Another nutrient which is sometimes un-proportional in aquaculture effluent is

phosphorous (P). A number of studies, such as Diem et al. (2017) or Monsees et al. (2017a),

found that total P accumulated over time in aquaponic systems growing lettuce. Excess P can

create antagonisms with micronutrients, such as Zn (Brunno da Silva Cerozi and Fitzsimmons,

2017). Alternatively, several other studies have found a lack of total P in their systems, well

below 17 mg/L in some cases (Endut et al. 2010; Goddek et al. 2015). This may be the result of

P precipitating as solid particles, and then becoming unavailable through the separation of solid

particles in filtration tanks. If there is a high level of Ca in the system water (common in areas

using water from basic limestone aquifers), P, specifically orthophosphate, will precipitate to

form calcium phosphate (Ca3(PO4)2) or dicalcium phosphate (CaHPO4) at a neutral pH (Rakocy

et al., 2006; Seawright et al., 1998). Ca3(PO4)2 is a solid which can be filtered out from the

system and is also not available for plant uptake (Rakocy, 2003). P deficiencies can lead to

stunted growth, specifically in heavier feeding crops such as fruits or vegetables (Brunno da

Silva Cerozi and Fitzsimmons, 2017). Despite significantly lower P concentrations, Blidariu, F.

C., Drasovean, A., & Grozea (2013) found that lettuce grown in aquaponic water containing P

concentrations of 3.2-3.6 mg/L had higher P tissue content than lettuce grow in field studies.

This may be a result of increased P availability in low P environments by PGPMs, such as

arbuscular mycorrhizae fungi as mentioned in section 3.1.6 (Akiyama et al., 2005; Bartelme et

al., 2018). Cerozi and Fitzsimmons (2017) found that P use efficiency was relatively high (71.7%

total, 42.3% in fish and 29.4% plants), in comparison to NUE.

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Despite aquaponic water samples being identified as having less nutrients in comparison

to standard hydroponic water (Bittsánszky et al., 2016), vegetative plants in aquaponics systems

have been reported to have similar or equal yields to those plants grown hydroponically. One

possible explanation for such a phenomenon, is that the nutrients in aquaponic water are in an

organic state, as opposed to a sterile ionic state found in most of the hydroponic water. This

organic source of nutrients may, A) reduce the measurement of nutrients actually in the water

(Rakocy, 2003), and B) promote growth by stimulating natural growing conditions as opposed to

sterile hydroponic conditions (Bӧhme, 1999; Goddek et al., 2015). These organic factors include

humic acids (fulvic acid) (Canellas et al. 2009; Haghighi et al. 2012), indole-3-acetic acid (IAA)

(Mangmang et al., 2015), rhizobacteria and arbuscular fungi (Bartelme et al. 2018). For example

Pantanella et al. (2012) and Delaide et al. (2016) found no difference in yields of lettuce

compared to lettuce grown in hydroponic systems, despite the aquaponic water of Delaide et al.

(2016) having 23% NO3-, 15% PO42-, 27% K+ and 8% Ca2+, of what was present in the

hydroponic water used in their study.

It is possible to manipulate the nutrient profile in the aquaculture effluent by changing the

diet fed. Seawright et al. (1998) were the first to investigate the nutrient profile created by

conventional fish feed, and the first to propose recommendations for feed manipulations to

provide better nutrients for plant production. They found that K, Mg, Mn, P, Na, and Zn were

nutrients which could be manipulated in the feed to increase plant productivity, while Fe and Cu

would not be able to be manipulated. A more recent study by Cerozi and Fitzsimmons (2017a)

investigated the use of a plant based feed with phytase. Plant proteins typically have P stored in

the form of phytate, which is unavailable for fish to assimilate. The addition of phytase to the

diet allows phytate to be broken down and assimilated in the fish. The study found that adding

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phytase in fish food increased growth in fish but lowered overall dissolved phosphorus in the

water. This, however, did not affect plant growth, indicating that a feed adaptation such as

phytase addition can increase P use efficiency (Brunnoda da Silva Cerozi and Fitzsimmons,

2017). Plant based fish feeds are a more sustainable form of feed and will most likely increase in

the aquaculture industry, considering that conventional fish feeds are primarily fish meal, which

is expensive, in shortage and an environmentally degrading feed component (Brunnoda da Silva

Cerozi and Fitzsimmons, 2017; Goddek et al., 2015). To the author’s knowledge, there is no

other comprehensive studies which investigate the effects of a custom fish feed in comparison to

a conventional fish feed in an aquaponic system. Future studies should investigate alternative

fish feeds, which produce effluent that is higher in K and Mg. In summary, suboptimal nutrient

concentrations compared to standard hydroponic solutions, appears to be a major limitation for

growing crops with higher nutrient requirements. Although there are several studies identifying

methods to improve crop yields through foliar sprays or adding fertilizer to the water, there is yet

to be research which investigates alternative methods of increasing nutrient concentrations in

aquaponic solution without using synthetic material, a fundamental pillar of sustainability in

aquaponics.

3.2.3 pH

The three main organisms in aquaponic systems: fish, nitrifying bacteria, and plants, have

different optimal pH ranges. Nile tilapia have an optimal pH between 7.0 and 9.0; the three

major nitrifying bacteria genera Nitrobacter, Nitrosomonas and Nitrospira have optimal pH

ranges of 7.5, 7.0-7.5 and 8.3, respectively (Antoniou et al., 1990; Goddek et al., 2015; Rakocy,

2003); and hydroponic plants perform optimally in a pH range of 5.8 to 6.2 (Cooper, 1979;

Sonneveld and Voogt, 2009). Given these obvious differences in optimal pHs, no one organism’s

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growth can be optimized without another’s being compromised, illustrating one of the major

challenges in any aquaponic system. Many aquaponic review articles have recommended an

optimal pH level, based on a compromise between optimal fish, bacteria and plant performance.

These optimal ranges vary depending on what aspect of aquaponics the author found to be more

valuable. For example, Rakocy et al. (2003), Tyson et al. (2008,) and Goddek et al. (2015)

suggested optimal pH ranges for aquaponics of 7.0, 6.8-7.0, and 7.5-8.0, respectively. Most

authors recommend a pH range that is more neutral, optimizing the nitrification process (pH

<7.0) over nutrient availability for plants. Alternatively, Tyson et al. (2008) used a lower pH of

6.0 to enhance nutrient uptake by plants; however, this resulted in increased fish mortality and a

significant decrease in nitrification. In mature aquaponic systems, water pH typically becomes

acidic over time. This is the result of acid (H+) production from the nitrification of ammonia,

which is continuously excreted from the gills of fish (Rakocy et al., 2004). Although most plants

in aquaponic systems release hydroxide –OH or bicarbonate HCO3- when assimilating nitrate, the

basic ions do not sufficiently counter the hydrogen released from nitrification (Goddek et al.,

2015). This acidic pH can be countered with the addition of KOH or Ca(OH) to the system

water, as commonly conducted in UVI systems (Rakocy, 2003). Although acidic pH levels may

be considered a challenge, Rakocy et al. (2003) outlined that if pH is remaining constant in a

mature system, it may be a result of excessive denitrification. If excess denitrification is

occurring, NUE will decrease along with available nitrogen for plant uptake. This also prevents

the addition of Ca(OH)2 or KOH to be added to the system, which may lead to deficiencies in Ca

or K. Alternatively, others have proposed the use of technology, such as a fluidized lime bed

reactor, to counter an acidic pH range (Goddek et al., 2015). A fluidized lime bed reactor would

supply a set quantity of dissolved limestone (CaCO3) directly into the system water which would

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produce hydroxide anions to increase pH. Although there are numerous publications on the

impact of pH in aquaponics, it appears as though it is not a major limiting factor if a

compensated range (6.7-7) is used to satisfy both plants and fish to some degree. Alternatively,

the use of DAS, as described in Section 2, is one solution to having varying optimal pH ranges.

Apart from nutritional related limitations, aquaponics is also burdened with the challenge of

handling solid accumulation in the system.

3.2.4 Solids

Build ups of organic material (un-eaten feed, algae, fungi, fish sludge) can be detrimental

to aquaponic systems. Although a small build-up of solid materials is necessary for providing a

constant concentration of nutrients through mineralization (via mineralization tanks or filtration

tanks), excessive solid build up can be harmful. Organic build ups or formation of sludge creates

anaerobic environments which can cause anaerobic bacteria to release carbon dioxide, ammonia,

hydrogen sulfide and methane, which are toxic to fish (Rakocy et al., 2006). Furthermore, if

solids build up around the roots of plants they can create an anaerobic environment and inhibit

oxygen to plant roots, which they require for nutrient uptake mechanisms (Rakocy et al., 2006).

Lowered dissolved oxygen (<7 mg/L) around the root zone also increases susceptibility to root

rot, caused by Pythium (Cherif et al., 1997). Anaerobic solid build ups also have the possibility

to create “off-flavours”. Off flavours are typically secondary metabolites, such as geosmin or 2-

methylisoborneol produced by Streptomyces, which can grow when there is an accumulation of

organic material containing phosphorous (Rurangwa and Verdegem, 2015). Although this is a

concern for some producers and consumers, a study investigating microbial communities in the

Wädenswil aquaponic facility found no Streptomyces present, nor was it reported in any of the

articles reviewed, indicating that this may not be a prevalent problem (Schmautz et al., 2017).

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Since aquaponic systems contain a high level of living microorganisms, it is possible that

pathogenic microbes or aquatic pests could develop, particularly if solids accumulate. One

example is the zooplankton pest. Zooplankton, specifically ostracods, pose a threat to aquaponic

systems by consuming root hairs, preventing nutrient uptake and inhibiting growth (Rakocy et

al., 2006). Andriani et al. (2018) found that the most common zooplankton in their aquaponic

systems were Brachionus sp. and Epistylis sp, the latter being a microorganism that causes

disease in fish. Another pest related organism that can develop from solid accumulation is

A.ocellatum; an “obligate ecto-parasitic, dinflagellate” which targets marine fish (Nozzi et al.

2016). Fortunatley, solid acumulation can be mitigated in most systems through sludge removing

clarifiers or filtration tanks which can be cleaned (Goddek et al., 2016; Rakocy et al., 2006). If a

disease or pest were to outbreak in an aquaponic system, there would be limited options to

remediate the situation, since pesticides and antibiotics cannot be used on plants/fish as their

effects are either unknown or harmful (Goddek et al., 2015). However, based on the literature

review conducted here, most studies investigating aquaponics have not reported any detrimental

disease out breaking in their system. It is possible that this is a result of beneficial microbial

communities living in the rhizosphere of plant roots which have antimicrobial properties,

allowing them to prevent disease (Gravel et al., 2015; Schmautz et al., 2017). Another concern is

the physical presence of microorganisms on marketable aquaponic produce, as their presence

may be high in the water. There is a number of restrictions on how many coliforms are allowed

in water, based on different global locations as described in Pantanella et al. (2015). A possible

method of decreasing these microbes is the use of UV light. Pantanella et al. (2015) found that

ultraviolet light decreased bacteria by a 1000 CFU/ml in the fish tanks, lowering the coliform

count below what was previously above the World Health Organization standard (1989). Few

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other studies have reported on the presence of microbes on marketable produce, and therefore

future research should be conducted on such matters. Since aquaponics produces food in close

proximity to fish waste, the safety and sterility of aquaponics produce is a concern. Aquaponics

lacks a regulating body to certify aquaponic products are truly aquaponically grown. A

certification system for aquaponic produce would greatly benefit the commercial industry as it

would ensure products are free of pathogens and safe for consumption, allow products to be

marketed at a higher price based on certification, and deter pseud-aquaponic cultivation (those

that use >50% synthetic fertilizer). This would further legitimize aquaponic produce for the

consumer, providing a guarantee that the product has been sustainably produced. Such factors

may increase the feasibility of aquaponic produce in the commercial market place and encourage

sustainable consumption.

4. Conclusion

Aquaponic systems carry great potential to overcome several sustainability challenges in

the agriculture sector; primarily the ability to produce high yields with minimal added nutrients,

while also greatly reducing nutrient discharge and water loss from aquaculture. The growing

interest in sustainable agriculture systems such as aquaponics, is illustrated in this review by the

exponential release of aquaponic publications in the last 3 years. From these recent

developments, clear trends, implications, and needs for future research on system design,

hydroponic components, optimal fish, plant and microorganism species have emerged, and are as

follows:

 It appears that a movement from CAS to DAS is primarily based on optimizing the

horticulture portion of aquaponics but lacks evidence that the change in systems will be

as economical and sustainably compromising, on account of increased required space and

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fertilizer for DAS. Given that commercial aquaponics is expanding rapidly, it is possible

that systems designs are rapidly adapted based on unproven premises, which may

endanger the sustainable effectiveness of these systems in the sector. This potential issue

can be clarified by well-designed experiments investigating the economics and

sustainability between CAS and DAS in a commercial context.

 Natural variations, management techniques, and application of hydroponic components in

aquaponic systems make it difficult to derive conclusions on which component is

optimal. For small scale and research aquaponic systems, a media bed hydroponic

component appears to be the most successful based on its ability to grow a variety of

plants and mitigate the use of a biofilter. For commercial aquaponic systems, a DWC

component appears to be optimal based on its low environmental impact, higher water

use efficiency, maximised root to water contact and consistent high plant yields.

 Tilapia, followed by Carp and African Catfish appear to be the most successful fish

species used in aquaponic systems. These fish are successful as a result of their ability to

handle high levels of nitrate and low oxygen. There has yet to be a study which

investigates the benefits of polyculture in aquaponic systems. Considering polyculture

may create a more optimally nutrient balanced solution for plant growth, future research

on this topic could have substantial value.

 Leafy greens such as varieties of lettuce, mint, and basil appear to be the most successful

plant species in aquaponics, as a result of their low nutrient requirements, universal

demand, and economical value in aquaponic systems. There are few studies referencing

the use of aquaponics for flowering crop production. Evaluating the use of aquaponics for

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a variety of flowering species, may allow the expansion of aquaponics into the

floriculture industry.

 It is possible Nitrospira has a major role in the nitrification process in aquaponics, over

past thought of nitrifiers such as Nitrobacter and Nitrosomonas. Future studies should

investigate the role of nitrifying bacteria in aquaponics through their NUE and

interactions with plant roots and surrounding microorganisms. Such studies may provide

information relating to improvements in nutrient extraction from aquaculture waste.

 PGPM in aquaponics are often suggested to be the reason plants can mitigate disease and

have equal yields to hydroponics, despite lower nutrient levels. There are few studies on

PGPM in aquaponic studies. Future PGPM studies on aquaponics will be imperative to

understanding the underlying mechanisms of nutrient dynamics in aquaponics systems,

which could be valuable for further improvement in all forms of agriculture. Given the

energy intensive requirements of fertilizer, the impact of growing plants with minimal

fertilizer can greatly reduce the carbon footprint of crops grown. This review suggests

that aquaponics is an ideal system to observe and understand the nature of PGPM, which

may allow for their application in other growing methods.

These trends and research suggestions can help both industry and academia establish what is

already successful in aquaponics and what research will be most influential. From a

horticulture perspective, the two following points appear to be the most limiting for the

industry and require future research:

 Aquaponics largest horticulture challenge is providing balanced nutrients for heavy

feeding plants. Specifically, aquaponic solution has been found to have insufficient K,

Mg, Ca and Fe for some plants. Several articles have found that these nutrient limitations

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can be overcome with nutrient additions to the system, or foliar sprays, however future

studies should investigate if nutrient levels in aquaponic water can be optimized through

custom fish feed or the use of polyculture, as such a solution would minimize the use of

synthetic fertilizers. Future studies are also needed to determine optimal Ca, Mg and Mn

additions for plants in aquaponic systems.

 There is a lack of standards to which aquaponic goods can be sold with ensured quality,

safety, and verification of true aquaponic cultivation. A clear set of aquaponic standards

should be developed to help create a more established market for aquaponic products.

References

Adler, P.R., Harper, J.K., Wade, E.M., Takeda, F., Summerfelt, S.T., 2000. Economic Analysis

of an Aquaponic System for the Integrated Production of Rainbow Trout and Plants. Int. J.

Recirc. Aquac. 1, 15–34. https://doi.org/10.21061/ijra.v1i1.1359

Akiyama, K., Matsuzaki, K.I., Hayashi, H., 2005. Plant sesquiterpenes induce hyphal branching

in arbuscular mycorrhizal fungi. Nature 435, 824–827. https://doi.org/10.1038/nature03608

Andriani, Y., Dhahiyat, Y., Zahidah, Subhan, U., Iskandar, Zidni, I., Mawardiani, T., 2018.

Effect of water irrigation volume on Capsicum frutescens growth and plankton abundance

in aquaponics system, in: IOP Conf. Series: Earth and Environmental Science. pp. 1–10.

https://doi.org/10.1088/1755-1315/139/1/012001

Antoniou, P., Hamilton, J., Koopman, B., Jain, R., Holloway, B., Lyberatos, G., Svoronos, S.A.,

1990. Effect of temperature and ph on the effective maximum specific growth rate of

nitrifying bacteria. Water Res. 24, 97–101. https://doi.org/10.1016/0043-1354(90)90070-M

Avis, T.J., Gravel, V., Antoun, H., Tweddell, R.J., 2008. Multifaceted beneficial effects of

39
 ACCEPTED MANUSCRIPT

rhizosphere microorganisms on plant health and productivity. Soil Biol. Biochem. 40,

1733–1740. https://doi.org/10.1016/j.soilbio.2008.02.013

Bailey, D.S., Ferrarezi, R.S., 2017. Valuation of vegetable crops produced in the UVI

Commercial Aquaponic System. Aquac. Reports 7, 77–82.

https://doi.org/10.1016/j.aqrep.2017.06.002

Bartelme, R.P., McLellan, S.L., Newton, R.J., 2017. Freshwater recirculating aquaculture system

operations drive biofilter bacterial community shifts around a stable nitrifying consortium

of ammonia-oxidizing archaea and comammox Nitrospira. Front. Microbiol. 8, 1–18.

https://doi.org/10.3389/fmicb.2017.00101

Bartelme, R.P., Oyserman, B.O., Blom, J.E., Sepulveda-Villet, O.J., Newton, R.J., 2018.

Stripping away the soil: Plant growth promoting microbiology opportunities in aquaponics.

Front. Microbiol. 9, 1–7. https://doi.org/10.3389/fmicb.2018.00008

Bittsánszky, A., Uzinger, N., Gyulai, G., Mathis, A., Junge, R., Villarroel, M., Kotzen, B.,

Kőmíves, T., 2016. Nutrient supply of plants in aquaponic systems. Ecocycles 2, 17–20.

https://doi.org/10.19040/ecocycles.v2i2.57

Blackburne, R., Vadivelu, V.M., Yuan, Z., Keller, J., 2007. Kinetic characterisation of an

enriched Nitrospira culture with comparison to Nitrobacter. Water Res. 41, 3033–3042.

https://doi.org/10.1016/j.watres.2007.01.043

Blidariu, F. C., Drasovean, A., & Grozea, A., 2013. Evaluation of phosphorus level in green

lettuce conventional grown under natural conditions and aquaponic system. Bull. UASVM

Anim. Sci. Biotechnol. 70, 128–135.

40
 ACCEPTED MANUSCRIPT

Bosma, R.H., Lacambra, L., Landstra, Y., Perini, C., Poulie, J., Schwaner, M.J., Yin, Y., 2017.

The financial feasibility of producing fish and vegetables through aquaponics. Aquac. Eng.

78, 146–154. https://doi.org/10.1016/j.aquaeng.2017.07.002

Buhmann, A.K., Waller, U., Wecker, B., Papenbrock, J., 2015. Optimization of culturing

conditions and selection of species for the use of halophytes as biofilter for nutrient-rich

saline water. Agric. Water Manag. 149, 102–114.

https://doi.org/10.1016/j.agwat.2014.11.001

Bӧhme, M., 1999. Effects of Lactate, Humate and Bacillus subtilis on the Growth of Tomato

PLants in Hydroponic Systems. Proc. Int. Sym. Grow. Media Hydroponics 481, 231–239.

Canellas, L.P., Spaccini, R., Piccolo, A., Dobbss, L.B., Okorokova-Façanha, A.L., Santos,

G.D.A., Olivares, F.L., Façanha, A.R., 2009. Relationships between chemical

characteristics and root growth promotion of humic acids isolated from Brazilian oxisols.

Soil Sci. 174, 611–620. https://doi.org/10.1097/SS.0b013e3181bf1e03

Canfield, D.E., Glazer, A.N., Falkowski, P.G., 2010. The Evolution and Future of Earth’s

Nitrogen Cycle. Science (80-. ). 330, 192–196. https://doi.org/10.1126/science.1186120

Cerozi, B. da S., Fitzsimmons, K., 2017. Phosphorus dynamics modeling and mass balance in an

aquaponics system. Agric. Syst. 153, 94–100. https://doi.org/10.1016/j.agsy.2017.01.020

Cerozi, B. da S., Fitzsimmons, K., 2017. Effect of dietary phytase on phosphorus use efficiency

and dynamics in aquaponics. Aquac. Int. 25, 1227–1238. https://doi.org/10.1007/s10499-

016-0109-7

Cerozi, B. da S., Fitzsimmons, K., 2016. Use of Bacillus spp. to enhance phosphorus availability

41
 ACCEPTED MANUSCRIPT

and serve as a plant growth promoter in aquaponics systems. Sci. Hortic. (Amsterdam). 211,

277–282. https://doi.org/10.1016/j.scienta.2016.09.005

Cherif, M., Tirilly, Y., Belanger, R.R., 1997. Effect of oxygen concentration on plant growth,

lipid peroxidation, and receptivity of tomato roots to Pythium under hydroponic conditions.

Eur. J. Plant Pathol. 103, 255–264.

Cooper, A., 1979. The ABC of NFT, 1st ed. Grower Books, London.

Daims, H., Lebedeva, E. V., Pjevac, P., Han, P., Herbold, C., Albertsen, M., Jehmlich, N.,

Palatinszky, M., Vierheilig, J., Bulaev, A., Kirkegaard, R.H., Von Bergen, M., Rattei, T.,

Bendinger, B., Nielsen, P.H., Wagner, M., 2015. Complete nitrification by Nitrospira

bacteria. Nature 528, 504–509. https://doi.org/10.1038/nature16461

Danaher, J.J., Shultz, R.C., Rakocy, J.E., Bailey, D.S., 2013. Alternative Solids Removal for

Warm Water Recirculating Raft Aquaponic Systems. J. World Aquac. Soc. 44, 374–383.

https://doi.org/10.1111/jwas.12040

Delaide, B., Goddek, S., Gott, J., Soyeurt, H., Jijakli, M.H., 2016. Lettuce (Lactuca sativa L. var.

Sucrine) growth performance in complemented aquaponic solution outperforms

hydroponics. Water (Switzerland) 8, 1–11. https://doi.org/10.3390/w8100467

Diem, T.N.T., Konnerup, D., Brix, H., 2017. Effects of recirculation rates on water quality and

Oreochromis niloticus growth in aquaponic systems. Aquac. Eng. 78, 95–104.

https://doi.org/10.1016/j.aquaeng.2017.05.002

Diver, S., 2006. Aquaponics — Integration of Hydroponics with Aquaculture. ATTRA Natl.

Sustain. Agric. Infromation Serv. https://doi.org/10.1146/annurev.genet.37.110801.143717

42
 ACCEPTED MANUSCRIPT

Eickhout, B., Bouwman, A.F., van Zeijts, H., 2006. The role of nitrogen in world food

production and environmental sustainability. Agric. Ecosyst. Environ. 116, 4–14.

https://doi.org/10.1016/j.agee.2006.03.009

El-Sayed, A.F.M., 2006. Tilapia Culture. CAB eBooks, Oceanography Department, Faculty of

Science, Alexandria University, Alexandria, Egypt.

https://doi.org/10.1079/9780851990149.0000

Endut, A., Jusoh, A., Ali, N., Wan Nik, W.B., Hassan, A., 2010. A study on the optimal

hydraulic loading rate and plant ratios in recirculation aquaponic system. Bioresour.

Technol. 101, 1511–1517. https://doi.org/10.1016/j.biortech.2009.09.040

Engle, C.R., 2015. Economics of Aquaponics. SRAC Publ. - South. Reg. Aquac. Cent. No. 5006,

1–4.

EPA, 2010. Methane and Nitrous Oxide Emissions from Natural Sources, U.S. Environmental

Protection Agency, Washington, DC, USA.

Fang, Y., Hu, Z., Zou, Y., Zhang, J., Zhu, Z., Zhang, J., Nie, L., 2017. Improving nitrogen

utilization efficiency of aquaponics by introducing algal-bacterial consortia. Bioresour.

Technol. 245, 358–364. https://doi.org/10.1016/j.biortech.2017.08.116

FAO, 2005. AQUASTAT Survey, Irrigation in Africa in Figures, in: FAO: Rome, Italy,.

Food and Agriculture Organization of the United Nations, 2018. The State of World Fisheries

and Aquaculture. Rome.

Food and Agriculture Organization of the United Nations, 1988. Definition of aquaculture, in:

Seventh Session of the IPFC Working Party of Expects on Aquaculture. pp. 1–3.

43
 ACCEPTED MANUSCRIPT

Forchino, A.A., Lourguioui, H., Brigolin, D., Pastres, R., 2017. Aquaponics and sustainability:

The comparison of two different aquaponic techniques using the Life Cycle Assessment

(LCA). Aquac. Eng. 77, 80–88. https://doi.org/10.1016/j.aquaeng.2017.03.002

Gajdanowicz, P., Michard, E., Sandmann, M., Rocha, M., Correa, L.G.G., Ramirez-Aguilar, S.J.,

Gomez-Porras, J.L., Gonzalez, W., Thibaud, J.-B., van Dongen, J.T., Dreyer, I., 2011.

Potassium (K+) gradients serve as a mobile energy source in plant vascular tissues. Proc.

Natl. Acad. Sci. 108, 864–869. https://doi.org/10.1073/pnas.1009777108

Gierth, M., Mäser, P., 2007. Potassium transporters in plants - Involvement in K+acquisition,

redistribution and homeostasis. FEBS Lett. 581, 2348–2356.

https://doi.org/10.1016/j.febslet.2007.03.035

Gilles, S., Ismiño, R., Sánchez, H., David, F., Núñez, J., Dugué, R., Darias, M.J., Römer, U.,

2014. An integrated closed system for fish-plankton aquaculture in Amazonian fresh water.

Animal 8, 1319–1328. https://doi.org/10.1017/S1751731114001165

Goda, A.M.A.-S., Essa, M.A., Hassaan, M.S., Sharawy, Z., 2015. Bio Economic Features for

Aquaponic Systems in Egypt. Turkish J. Fish. Aquat. Sci. 15, 525–532.

https://doi.org/10.4194/1303-2712-v15

Goddek, S., Delaide, B., Mankasingh, U., Ragnarsdottir, K.V., Jijakli, H., Thorarinsdottir, R.,

2015. Challenges of sustainable and commercial aquaponics. Sustain. 7, 4199–4224.

https://doi.org/10.3390/su7044199

Goddek, S., Keesman, K.J., 2018. The necessity of desalination technology for designing and

sizing multi-loop aquaponics systems. Desalination 428, 76–85.

https://doi.org/10.1016/j.desal.2017.11.024

44
 ACCEPTED MANUSCRIPT

Goddek, S., Schmautz, Z., Scott, B., Delaide, B., Keesman, K., Wuertz, S., Junge, R., 2016. The

Effect of Anaerobic and Aerobic Fish Sludge Supernatant on Hydroponic Lettuce.

Agronomy 6, 1–12. https://doi.org/10.3390/agronomy6020037

Graber, A., Junge, R., 2009. Aquaponic Systems: Nutrient recycling from fish wastewater by

vegetable production. Desalination 246, 147–156.

https://doi.org/10.1016/j.desal.2008.03.048

Gravel, V., Dorais, M., Dey, D., Vandenberg, G., 2015. Fish effluents promote root growth and

suppress fungal diseases in tomato transplants. Can. J. Plant Sci. 95, 427–436.

https://doi.org/10.4141/cjps-2014-315

Gullian Klanian, M., Delgadillo Diaz, M., Aranda, J., Rosales Juárez, C., 2018. Integrated effect

of nutrients from a recirculation aquaponic system and foliar nutrition on the yield of

tomatoes Solanum lycopersicum L. and Solanum pimpinellifolium. Environ. Sci. Pollut.

Res. https://doi.org/10.1007/s11356-018-1817-5

Haghighi, M., Kafi, M., Fang, P., 2012. Photosynthetic Activity and N Metabolism of Lettuce as

Affected by Humic Acid. Int. J. Veg. Sci. 18, 182–189.

https://doi.org/10.1080/19315260.2011.605826

Hargreaves, J.A., 1998. Nitrogen biogeochemistry of aquaculture ponds. Aquaculture 166, 181–

212. https://doi.org/10.1016/S0044-8486(98)00298-1

Hu, Z., Lee, J.W., Chandran, K., Kim, S., Brotto, A.C., Khanal, S.K., 2015. Effect of plant

species on nitrogen recovery in aquaponics. Bioresour. Technol. 188, 92–98.

https://doi.org/10.1016/j.biortech.2015.01.013

45
 ACCEPTED MANUSCRIPT

IAFFD, 2018. Feed Ingredient Composition Database [WWW Document]. URL

http://www.iaffd.com/home.html?v=4.01

Jiang, Q., Dilixiati, A., Zhang, W., Li, W., Wang, Q., Zhao, Y., Yang, J., Li, Z., 2014. Effect of

nitrite exposure on metabolic response in the freshwater prawn Macrobrachium nipponense.

Cent. Eur. J. Biol. 9, 86–91. https://doi.org/10.2478/s11535-013-0167-4

Junge, R., König, B., Villarroel, M., Komives, T., Jijakli, M.H., 2017. Strategic points in

aquaponics. Water (Switzerland) 9, 1–9. https://doi.org/10.3390/w9030182

Khandaker, M., Kotzen, B., 2018. The potential for combining living wall and vertical farming

systems with aquaponics with special emphasis on substrates. Aquac. Res. 49, 1454–1468.

https://doi.org/10.1111/are.13601

Kloas, W., Groß, R., Baganz, D., Graupner, J., Monsees, H., Schmidt, U., Staaks, G., Suhl, J.,

Tschirner, M., Wittstock, B., Wuertz, S., Zikova, A., Rennert, B., 2015. A new concept for

aquaponic systems to improve sustainability, increase productivity, and reduce

environmental impacts. Aquac. Environ. Interact. 7, 179–192.

https://doi.org/10.3354/aei00146

Knaus, U., Palm, H.W., 2017. Effects of the fish species choice on vegetables in aquaponics

under spring-summer conditions in northern Germany (Mecklenburg Western Pomerania).

Aquaculture 473, 62–73. https://doi.org/10.1016/j.aquaculture.2017.01.020

Kong, Y., Zheng, Y., 2014. Potential of producing Salicornia bigelovii hydroponically as a

vegetable at moderate NaCl salinity. HortScience 49, 1154–1157.

https://doi.org/10.1139/b03-086

46
 ACCEPTED MANUSCRIPT

König, B., Janker, J., Reinhardt, T., Villarroel, M., Junge, R., 2018. Analysis of aquaponics as an

emerging technological innovation system. J. Clean. Prod. 180, 232–243.

https://doi.org/10.1016/j.jclepro.2018.01.037

Krom, M.D., Ellner, S., van Rijn, J., Neori, A., 1995. Nitrogen and phosphorus cycling and

transformations in a prototype “non-polluting” integrated mariculture system, Eilat, Israel.

Mar. Ecol. Prog. Ser. 118, 25–36. https://doi.org/10.3354/meps118025

Lennard, W.A., 2015. AQUAPONICS: a nutrient dynamic process and the relationship to fish

feeds. J. World Aquac. Soc. September, 20–23.

Lennard, W.A., Leonard, B. V., 2006. A comparison of three different hydroponic sub-systems

(gravel bed, floating and nutrient film technique) in an Aquaponic test system. Aquac. Int.

14, 539–550. https://doi.org/10.1007/s10499-006-9053-2

Liang, J.Y., Chien, Y.H., 2013. Effects of feeding frequency and photoperiod on water quality

and crop production in a tilapia-water spinach raft aquaponics system. Int. Biodeterior.

Biodegrad. 85, 693–700. https://doi.org/10.1016/j.ibiod.2013.03.029

Love, D.C., Fry, J.P., Genello, L., Hill, E.S., Frederick, J.A., Li, X., Semmens, K., 2014. An

international survey of aquaponics practitioners. PLoS One 9, 1–10.

https://doi.org/10.1371/journal.pone.0102662

Love, D.C., Fry, J.P., Li, X., Hill, E.S., Genello, L., Semmens, K., Thompson, R.E., 2015a.

Commercial aquaponics production and profitability: Findings from an international survey.

Aquaculture 435, 67–74. https://doi.org/10.1016/j.aquaculture.2014.09.023

Love, D.C., Uhl, M.S., Genello, L., 2015b. Energy and water use of a small-scale raft aquaponics

47
 ACCEPTED MANUSCRIPT

system in Baltimore, Maryland, United States. Aquac. Eng. 68, 19–27.

https://doi.org/10.1016/j.aquaeng.2015.07.003

Mangmang, J.S., Deaker, R., Rogers, G., 2015. Response of lettuce seedlings fertilized with fish

effluent to Azospirillum brasilense inoculation. Biol. Agric. Hortic. 31, 61–71.

https://doi.org/10.1080/01448765.2014.972982

Marques, B., Calado, R., Lillebø, A.I., 2017. New species for the biomitigation of a super-

intensive marine fish farm effluent: Combined use of polychaete-assisted sand filters and

halophyte aquaponics. Sci. Total Environ. 599–600, 1922–1928.

https://doi.org/10.1016/j.scitotenv.2017.05.121

Mashifane, T.B., Moyo, N.A.G., 2014. Acute toxicity of selected heavy metals to Oreochromis

mossambicus fry and fingerlings. African J. Aquat. Sci. 39, 279–285.

https://doi.org/10.2989/16085914.2014.960358

Maucieri, C., Nicoletto, C., Junge, R., Schmautz, Z., Sambo, P., Borin, M., 2018. Hydroponic

systems and water management in aquaponics: A review. Ital. J. Agron. 13, 1–11.

https://doi.org/10.4081/ija.2017.1012

Mcmurtry, M.R., Sanders, D.C., Cure, J.D., Hudson, R.G., Haning, B.C., Amand, P.C.S., 1997.

Efficiency of water use of an integrated fish/vegetable co-culture system. J. World Aquac.

Soc. 28, 420–428. https://doi.org/10.1111/j.1749-7345.1997.tb00290.x

Miličić, V., Thorarinsdottir, R., Dos Santos, M., Hančič, M.T., 2017. Commercial aquaponics

approaching the European market: To consumers’ perceptions of aquaponics products in

Europe. Water (Switzerland) 9, 1–22. https://doi.org/10.3390/w9020080

48
 ACCEPTED MANUSCRIPT

Molovan, I., Băla, M., 2015. Analysis of aquaponic organic hydroponics from the perspective of

setting costs and of maintenance on substratum and floating shelves systems. J. Hortic. For.

Biotechnol. 19, 73–76.

Monsees, H., Keitel, J., Paul, M., Kloas, W., Wuertz, S., 2017a. Potential of aquacultural sludge

treatment for aquaponics: Evaluation of nutrient mobilization under aerobic and anaerobic

conditions. Aquac. Environ. Interact. 9, 9–18. https://doi.org/10.3354/aei00205

Monsees, H., Kloas, W., Wuertz, S., 2017b. Decoupled systems on trial: Eliminating bottlenecks

to improve aquaponic processes. PLoS One 12, 1–18.

https://doi.org/10.1371/journal.pone.0183056

Mudahar, M.S., Hignett, T.P., 1985. Energy Efficiency in Nitrogen Fertilizer Production. Energy

Agric. Elsevier Sci. Publ. B.V 4, 159–177. https://doi.org/10.1016/0167-5826(85)90014-2

Mylonas, V.A., McCants, C.B., 1980. Effects of humic and fulvic acids on growth of tobacco.

Plant Soil 54, 485–490. https://doi.org/10.1007/BF02181841

Naegel, L.C.A., 1977. Combined production of fish and plants in recirculating water.

Aquaculture 10, 17–24. https://doi.org/10.1016/0044-8486(77)90029-1

Nozzi, V., Strofaldi, S., Piquer, I.F., Di Crescenzo, D., Olivotto, I., Carnevali, O., 2016.

Amyloodinum ocellatum in Dicentrarchus labrax: Study of infection in salt water and

freshwater aquaponics. Fish Shellfish Immunol. 57, 179–185.

https://doi.org/10.1016/j.fsi.2016.07.036

Oelkers, E.H., Valsami-Jones, E., 2008. Phosphate Mineral Reactivity and Global Sustainability.

Elements 4, 83–87.

49
 ACCEPTED MANUSCRIPT

Palm, H.W., Knaus, U., Appelbaum, S., Goddek, S., Strauch, S.M., Vermeulen, T., Haїssam

Jijakli, M., Kotzen, B., 2018. Towards commercial aquaponics: a review of systems,

designs, scales and nomenclature. Aquac. Int. 1–30. https://doi.org/10.1007/s10499-018-

0249-z

Palm, H.W., Seidemann, R., Wehofsky, S., Knaus, U., 2014. Significant factors affecting the

economic sustainability of closed aquaponic systems. Part I: System design, chemo-physical

parameters and general aspects. AACL Bioflux 7, 20–32.

Pantanella, E., Cardarelli, M., Colla, G., Rea, E., Marcucci, A., 2012. Aquaponics vs.

Hydroponics: Production and Quality of Lettuce Crop. Acta Hortic. 927, 887–894.

https://doi.org/10.17660/ActaHortic.2012.927.109

Pantanella, E., Cardarelli, M., Di Mattia, E., Colla, G., 2015. Aquaponics and food safety:

Effects of UV sterilization on total coliforms and lettuce production. Acta Hortic. 1062, 71–

76. https://doi.org/10.17660/ActaHortic.2015.1062.8

Pattillo, D.A., 2017. An Overview of Aquaponic Systems: Hydroponic Components. NRAC

Tech. Bull. Ser. 19, 1–10.

Radzki, W., Gutierrez Mañero, F.J., Algar, E., Lucas García, J.A., García-Villaraco, A., Ramos

Solano, B., 2013. Bacterial siderophores efficiently provide iron to iron-starved tomato

plants in hydroponics culture. Antonie van Leeuwenhoek, Int. J. Gen. Mol. Microbiol. 104,

321–330. https://doi.org/10.1007/s10482-013-9954-9

Rakocy, J.E., 2012. Chapter 14: aquaponics–integrating fish and plant culture., in: Aquaculture

Production System. Wiley, Hoboken, pp. 343–386.

50
 ACCEPTED MANUSCRIPT

Rakocy, J.E., 2003. Aquaponics — Integrating Fish and Plant Culture, in: Aquaculture

Production Systems. pp. 343–386. https://doi.org/10.1002/9781118250105.ch14

Rakocy, J.E., Bailey, D.S., Shultz, R.C., Thoman, E.S., 2010. Update on tilapia and vegetable

production in the UVI aquaponic system, in: Proceedings from the 6th International

Symposium on Tilapia in Aquaculture. pp. 1–15.

Rakocy, J.E., Masser, M.P., Losordo, T.M., 2006. Recirculating aquaculture tank production

systems: Aquaponics- integrating fish and plant culture. SRAC Publ. - South. Reg. Aquac.

Cent. 16. https://doi.org/454

Rakocy, J.E., Shultz, R.C., Bailey, D.S., Thoman, E.S., 2004. Aquaponic production of tilapia

and basil: Comparing a batch and staggered cropping system. Acta Hortic. 648, 63–69.

https://doi.org/10.17660/ActaHortic.2004.648.8

Resh, H.M., 2012. Hydroponic Food Production: A Definitive Guidebook for the Advanced

Home Gardener and the Commercial Hydroponic Grower. CRC Press, Boca Raton, FL,

USA.

Resh, H.M., 1995. Hydroponic food production, Woodbridge Press Publishing Company,. Santa

Barbara, California.

Roosta, H.R., 2014. Effects of Foliar Spray of K on Mint, Radish, Parsley and Coriander Plants

in Aquaponic System. J. Plant Nutr. 37, 2236–2254.

https://doi.org/10.1080/01904167.2014.920385

Roosta, H.R., Mohsenian, Y., 2012. Effects of foliar spray of different Fe sources on pepper

(Capsicum annum L.) plants in aquaponic system. Sci. Hortic. (Amsterdam). 146, 182–191.

51
 ACCEPTED MANUSCRIPT

https://doi.org/10.1016/j.scienta.2012.08.018

Rose, A.L., Waite, T.D., 2002. Kinetic model for FE(II) oxidation in seawater in the absence and

presence of natural organic matter. Environ. Sci. Technol. 36, 433–444.

https://doi.org/10.1021/es0109242

Ru, D., Liu, J., Hu, Z., Zou, Y., Jiang, L., Cheng, X., Lv, Z., 2017. Improvement of aquaponic

performance through micro- and macro-nutrient addition. Environ. Sci. Pollut. Res. 24,

16328–16335. https://doi.org/10.1007/s11356-017-9273-1

Rupasinghe, J.W., Kennedy, J.O.S., 2010. Economic benefits of integrating a hydroponic-lettuce

system into a barramundi fish production system. Aquac. Econ. Manag. 14, 81–96.

https://doi.org/10.1080/13657301003776631

Rurangwa, E., Verdegem, M.C.J., 2015. Microorganisms in recirculating aquaculture systems

and their management. Rev. Aquac. 7, 117–130. https://doi.org/10.1111/raq.12057

Savidov, N. a., Hutchings, E., Rakocy, J.E., 2007. Fish and Plant Production in a Recirculating

Aquaponic System a New Approach to Sustainable Agriculture in Canada. Acta Hortic.

742, 209–222. https://doi.org/10.17660/ActaHortic.2007.742.28

Schmautz, Z., Graber, A., Jaenicke, S., Goesmann, A., Junge, R., Smits, T.H.M., 2017. Microbial

diversity in different compartments of an aquaponics system. Arch. Microbiol. 199, 613–

620. https://doi.org/10.1007/s00203-016-1334-1

Schmautz, Z., Loeu, F., Liebisch, F., Graber, A., Mathis, A., Bulc, T.G., Junge, R., 2016. Tomato

productivity and quality in aquaponics: Comparison of three hydroponic methods. Water

(Switzerland) 8, 1–22. https://doi.org/10.3390/w8110533

52
 ACCEPTED MANUSCRIPT

Seawright, D.E., Stickney, R.R., Walker, R.B., 1998. Nutrient dynamics in integrated

aquaculture-hydroponics systems. Aquaculture 160, 215–237.

https://doi.org/10.1016/S0044-8486(97)00168-3

Shiklomanov, I.., 1998. World water resources A new appraisal and assessment for the 21st

century, International Hydrological Programme.

Sikawa, D.C., Yakupitiyage, A., 2010. The hydroponic production of lettuce (Lactuca sativa L)

by using hybrid catfish (Clarias macrocephalus × C. gariepinus) pond water: Potentials and

constraints. Agric. Water Manag. 97, 1317–1325.

https://doi.org/10.1016/j.agwat.2010.03.013

Silva, L., Valdés-Lozano, D., Escalante, E., Gasca-Leyva, E., 2018. Dynamic root floating

technique: An option to reduce electric power consumption in aquaponic systems. J. Clean.

Prod. 183, 132–142. https://doi.org/10.1016/j.jclepro.2018.02.086

Sonneveld, C., Voogt, W., 2009. Nutrient Management in Substrate Systems, in: Plant Nutrition

of Greenhouse Crops. Springer, Heidelberg, The Netherlands, pp. 277–312.

https://doi.org/10.1007/978-90-481-2532-6

Timmons, M.., Ebling, J.., 2010. Recirculating Aquaculture, 2nd ed. Ithaca NY, USA.

Tokunaga, K., Tamaru, C., Ako, H., Leung, P., 2015. Economics of small-scale commercial

aquaponics in hawai’i. J. World Aquac. Soc. 46, 20–32. https://doi.org/10.1111/jwas.12173

Turcios, A.E., Papenbrock, J., 2014. Sustainable treatment of aquaculture effluents-What can we

learn from the past for the future? Sustain. 6, 836–856. https://doi.org/10.3390/su6020836

Tyson, R. V., Simonne, E.H., Treadwell, D.D., White, J.M., Simonne, A., 2008. Reconciling pH

53
 ACCEPTED MANUSCRIPT

for ammonia biofiltration and cucumber yield in a recirculating aquaponic system with

perlite biofilters. HortScience 43, 719–724.

Tyson, R. V., Treadwel, D.D., Simonne, E.H., 2011. Opportunities and challenges to

sustainability in aquaponic systems. Horttechnology 21, 6–13.

Vermeulen, T., Kamstra, A., 2013. The need for systems design for robust aquaponic systems in

the urban environment, in: IS on Soilless Cultivation. pp. 71–78. https://doi.org/DOI:

10.17660/ActaHortic.2013.1004.6

Villarroel, M., Alvariño, J.M.R., Duran, J.M., 2011. Aquaponics: integrating fish feeding rates

and ion waste production for strawberry hydroponics. Spanish J. Agric. Res. 9, 537.

https://doi.org/10.5424/sjar/20110902-181-10

Wongkiew, S., Hu, Z., Chandran, K., Lee, J.W., Khanal, S.K., 2017a. Nitrogen transformations

in aquaponic systems: A review. Aquac. Eng. 76, 9–19.

https://doi.org/10.1016/j.aquaeng.2017.01.004

Wongkiew, S., Popp, B.N., Kim, H.J., Khanal, S.K., 2017b. Fate of nitrogen in floating-raft

aquaponic systems using natural abundance nitrogen isotopic compositions. Int. Biodeterior.

Biodegrad. 125, 24–32. https://doi.org/10.1016/j.ibiod.2017.08.006

Zou, Y., Hu, Z., Zhang, J., Xie, H., Guimbaud, C., Fang, Y., 2016a. Effects of pH on nitrogen

transformations in media-based aquaponics. Bioresour. Technol. 210, 81–87.

https://doi.org/10.1016/j.biortech.2015.12.079

Zou, Y., Hu, Z., Zhang, J., Xie, H., Liang, S., Wang, J., Yan, R., 2016b. Attempts to improve

nitrogen utilization efficiency of aquaponics through nitrifies addition and filler gradation.

54
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Environ. Sci. Pollut. Res. 23, 6671–6679. https://doi.org/10.1007/s11356-015-5898-0

55
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 Publications on aquaponics have increased exponentially in the past 3 years

 Aquaponic systems are moving from coupled to decoupled systems
 Tilapia paired with dark leafy greens are the commonly used species in aquaponics
 Plant growth promoting microbes play a vital role in the success of aquaponics
 The most prominent horticultural challenge in aquaponics is suboptimal nutrients