Aquaculture 466 (2017) 51–63

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Aquaculture

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Effects of fish biology on ebb and flow aquaponical cultured herbs in

northern Germany (Mecklenburg Western Pomerania)
U. Knaus ⁎, H.W. Palm
University of Rostock, Faculty of Agricultural and Environmental Sciences, Department of Aquaculture and Sea-Ranching, Justus-von-Liebig-Weg 6, D-18059 Rostock, Germany

a r t i c l e i n f o a b s t r a c t

Article history: Two identical coupled warmwater ebb-and-flood gravel aquaponic systems (unit I, unit II) without additional
Received 4 December 2015 biofilter were evaluated for the effect of fish species choice onto culinary herbs (basil - Ocimum basilicum, parsley
Received in revised form 18 July 2016 - Petroselinum crispum and marjoram - Origanum majorana) and resulting water parameters. The daily feed input
Accepted 12 September 2016
increased over the experiment (70–450 g, total 70 days), resulting in an maximum feed input of 150 g d−1, or 25%
Available online 14 September 2016
less than during an earlier study under the presence of a 60 L trickling biofilter. Using dissolved oxygen to indicate
Keywords:
system performance, the experiment had a run in phase, exponential phase, and accumulation phase. Lowest dis-
Aquaponics solved oxygen (DO) concentrations were measured inside the hydroponic units, with no general differences in
Ebb-and-flood chemo-physical parameters between each separate plant box. Nile tilapia red strain fingerlings (Oreochromis
Nile tilapia niloticus; initial weight 3.32 g ± 1.63) were stocked in aquaponic unit I and African catfish fingerlings (Clarias
African catfish gariepinus; initial weight 3.92 g ± 1.63) in unit II. Due to the high feed input at the beginning of the experiments,
Air breathing specific growth rate and feed conversion ratio were very good, and significantly better for C. gariepinus (3.30%
Synergy d−1 ± 0.02, 0.61 ± 0.01) compared with O. niloticus (2.81% d−1 ± 0.12, 0.91 ± 0.08). Feed conversion
Herbs
corresponded with much younger fish with effective feeding under restricted food input. The oxygen consump-
tion was lower for C. gariepinus with a difference of 8.34% to O. niloticus. Due to reduced natural light illumination
and water temperature (winter season), plant growth was generally reduced but two times better in the O.
niloticus stocked unit (0.30 g ± 0.37 in basil, 0.55 g ± 0.39 in parsley) compared with C. gariepinus (0.12 g ±
0.21 in basil, 0.27 g ± 0.31 in parsley). A newly calculated aquaponic growth factor (AGF) illustrates the opposite
fish and plant yields for both fish species.
Statement of relevance: The influence of different fish species on plant growth has been examined only rarely in
aquaponics. We hereby describe the influence of different types of fish on physical parameters and plant growth
in coupled aquaponic systems for diversification.
© 2016 Elsevier B.V. All rights reserved.

1. Introduction
Aquaponics, the integration of aquaculture and hydroponics, can
combine a wide range of suitable fish and plants. The composition of
the fish feed and the respective fish physiology are the main factors af-
fecting nutrient availability inside the system. So far, species of
Perciformes in particular “Tilapia” are the mostly frequently used fish
in aquaponics (Love et al., 2015; Rakocy et al., 2006). Blue tilapia
(Sarotherodon aurea) was used by Watten & Busch (1984), hybrid tilapia
(Oreochromis mossambicus × Oreochromis niloticus) by McMurtry et al.
(1997), Nile tilapia ‛red strain’ hybrid (Oreochromis niloticus × O. aureus)
by Kotzen & Appelbaum (2010) or the red tilapia (Oreochromis sp.) by
Liang & Chien (2013). Most often, Nile tilapia (Oreochromis niloticus)
⁎ Corresponding author.
E-mail addresses: ulrich.knaus@uni-rostock.de (U. Knaus), harry.palm@uni-rostock.de
(H.W. Palm).
was used, e.g. by Rakocy (1989a), Seawright et al. (1998), Essa et al.
(2008), Graber & Junge (2009), Rennert et al. (2011), Simeonidou et al.
(2012) and Palm et al. (2014a). In contrast, members of Siluriformes
such as African catfish (Clarias gariepinus) were newly investigated by
Endut et al. (2009) and Palm et al. (2014b) and hybrid catfish (Clarias
macrocephalus × C. gariepinus) by Sikawa & Yakupitiyage (2010). These
studies demonstrated the outstanding importance of Perciformes and
Siluriformes for aquaponical production systems.
Culinary herbs are the best choice for aquaponical plant production
with the highest level of income per unit of culture area per unit time
(Rakocy, 2012). According to Love et al. (2015), basil (Ocimum basilicum)
was the most frequently used crop under commercial aquaponic produc-
tion. Basil production was studied in an outdoor commercial-scale raft
aquaponic system in the Caribbean (University of the Virgin Islands,
USA) with a mean yield of 2.0 kg m−2 in batch production (Rakocy
et al., 2003) or an annual projected basil yield of 25.0 kg m−3 (Rakocy
et al., 2004). Kotzen & Appelbaum (2010) assessed qualitative and quan-
titative growth parameters of basil and other herbs in brackish and

http://dx.doi.org/10.1016/j.aquaculture.2016.09.025
0044-8486/© 2016 Elsevier B.V. All rights reserved.
52 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63

freshwater aquaponic systems under desert conditions of the Negev. Not low-tech warmwater, coupled ebb-and-flood aquaponic units with
all plant species showed good growth, which indicated benefits of selec- the use of gravel substrate adopted from Palm et al. (2014a,b) were
tive plant species cultivation. Only moderate growth was observed in tested in a medium temperate glasshouse during wintertime condi-
basil (Ocimum basilicum) and purple basil (Ocimum basilicum tions in the northern hemisphere. Cultivated plant species were
‘purparescens’) in contrast to better growth of parsley (Petroselinum basil (Ocimum basilicum), parsley (Petroselinum crispum) and marjo-
sp.) in freshwater and partly in brackish water. Campos Pulido (2013) ram (Origanum majorana). The experimental design included an in-
found reduced growth of Petroselinum crispum with higher salinities creasing feed input to identify the changing water parameters
and effluents from Oreochromis niloticus × O. aureus production. inside the sampled system. The effects of the omission of the
Danaher (2013) showed no differences of plant growth parameters in biofilter, extension of the plant units (20%) and addition of a drip-
parsley (Petroselinum crispum) between an inorganic fertilisation and a water overhead distribution onto (I) the maximum possible daily
leachate from a geotextile bag dewatering with aquaculture effluents feed input, (II) water parameters, (III) plant growth, and (IV) system
from Nile tilapia (Oreochromis niloticus). Buzby et al. (2016) reported productivity are discussed.
no different growth of P. crispum with the use of effluents from rainbow
trout (Oncorhynchus mykiss) under a low, high and amended high flow
2. Materials and methods
treatment. Marjoram (Origanum majorana) was rarely investigated
under aquaponical production. Hundley (2013) reported favourable
2.1. System and experimental design
growth of O. majorana in combination with O. niloticus in Brasilia. Thus,
O. basilicum has a wider importance for aquaponical production, while
Two identical coupled ebb-and-flood substrate aquaponic units
P. crispum and O. majorana are newly introduced in integrated produc-
(Fig. 1) in a temperate glasshouse with limed windows (50 m² operat-
tion systems.
ing area for both systems) were stocked with the Nile tilapia
In coupled aquaponics fish species selection and the choice of culti-
(Oreochromis niloticus, unit I) and African catfish (Clarias gariepinus,
vated plants may influence system stability parameters. However, in-
unit II). Each aquaponics contained a fibreglass fish tank (2.05 ×
vestigations on effects of different fish species or Crustacea onto the
2.05 × 0.93 m, AquaLogistik, Möhnesee-Wippringsen, Germany) filled
same plants are rare. Sace & Fitzsimmons (2013) found better growth
with appr. 1800.00 L freshwater, a clarifier (IBC) with 800 L, five plant
of vegetables with freshwater prawn (Macrobrachium rosenbergii) and
boxes (1.00 × 2.00 × 0.30 m) and a sump (0.61 m³), each system with
Nile tilapia (O. niloticus) reared together in polyculture in a coupled
a total volume of 3.81 m³. Mean water exchange rate was 1.48% and
raft aquaponic system. With prawn the highest yield was observed in
mean water removal was 0.26% (n = 18 days). The water was circulated
pac choi (Brassica rapa) followed by Chinese cabbage (Brassica rapa
with a single pump (UP150, 9000 L h−1, 160 W, Seerose Pumpentechnik
pekinensis) and lettuce (Lactuca sativa). Water quality parameters
Turkowski GmbH, Germany), connected to a single flow-type heater
showed better values for plant growth in combination with
(3.00 KW) and automatic temperature control at 25 °C. A linear forcer
M. rosenbergii with a higher aquaponic system stability. A most recent
air pump LA-120A (118 W, Nitto Kohki GmbH, Germany) with 4 air dif-
study with different fish species (C. gariepinus, O. niloticus) and basil
fusers in the fish tank provided adequate aeration. Each of the five
(Ocimum basilicum), lettuce (Lactuca sativa), cucumber (Cucumis
wooden plant boxes contained a polyethylene foil (1.50 mm) and
sativus) and tomato (Solanum lycopersicum) by Palm et al. (2014b)
400 kg gravel substrate (grain size of 16–32 mm, coarse gravel, pebble)
demonstrated better growth of plants with significant greater yield of
with a maximum water level of 20 cm, resulting in a maximum water
lettuce and cucumber fruits in combination with O. niloticus. The yield
volume of 120 L per box (measured). The ratio of the fish tank
of basil showed a better mean output of 159.00 g (± 96.91) with the
(1800 L), clarifier (800 L), hydroponic unit (600 L) and sump (610 L)
Nile tilapia in contrast to 117.94 g (± 92.30) with the African catfish.
was 2.25:1:0.75:0.76 (fish tank:hydroponic unit = 3.00).
These studies demonstrated the influence of fish species and Crustacea
The five plant boxes of each aquaponic unit were equipped with a
choice on plant growth and system stability parameters. For the under-
decentralised overhead drip irrigation system 30 cm above the gravel
standing of aquaponics biotechnology and the diversification of com-
substrate, filled with nutrient enriched water from the main water
bined fish and plant production further studies are necessary.
pipe (Fig. 1). The process water dripped more uniform onto the whole
The aquaponic set-up used in the present study was adopted from
plant box surface area. The overhead drip system consisted of three
Palm et al. (2014a) with two identical coupled small-scale aquaponic
polyvinylchloride pipes with 14 water outlets (Fig. 2B). The water in-
units with an 8 m² gravel substrate hydroponics and a total water vol-
flow of each plant box was adjusted to 2 L min−1. The outlets of the
ume of 3.75 m³. Dissolved oxygen was identified as the most important
plant boxes were constructed as a bell siphon (Bruno et al., 2011),
water parameter which was used as an indicator of system stability due
with an average flooding interval of 17.58 min (± 4.04) and a mean
to lower oxygen levels as a result of higher nutrient loads during the
water outflow time of 9.46 min (± 2.76), resulting in two ebb-and-
production period. Palm et al. (2014a) subdivided their experiment of
flood intervals per hour. For data collection of physical and chemical
160 days into three sub-experiments (SE I: 49 days: “run in phase”, SE
water parameters three plant boxes (boxes I, III, V) of each aquaponic
II: 56 days: “exponential phase” and SE III: 55 days: “steady phase”), de-
unit were equipped with three parallel rows of PVC test tubes, situat-
pending on the oxygen curves under cultivation of small sized O.
ed at the front (n = 3) of each box, the middle (n = 3) and the back
niloticus and different plants like tomato, basil or aubergine. During
position (n = 3) close to the outlet (Figs. 1, 2B).
their experiment, the feed input constantly increased (run-in and expo-
nential phase) until the oxygen level dropped significantly, demonstrat-
ing a system nutrient overload. The reduced feed input level of 200 g per 2.2. Fish and plant species
day resulted in a balanced production with a relatively stable oxygen
level (steady phase). The aquaponics from the present experiment dif- Three weeks before the experiment during winter season
fered from Palm et al. (2014a) in simple system changes, with the exclu- (13.11.2013–21.01.2014, total of 70 days), Nile tilapia red strain
sion of a 60 L biofilter, the increase of 20% hydroponic surface (from 8 to (Oreochromis niloticus) all male population (batch, 554 fish m−3) origi-
10 m2, addition of one plant box) with a total water volume of 3.81 m³ nating from Kirschauer Aquakulturen GmbH (Germany) were stocked
and a better irrigation system for the nutrient enriched water into the into the cleaned aquaponics of unit I. Two weeks later mixed sex African
hydroponics. catfish (Clarias gariepinus, batch, 347 fish m−3) originating from a local
The main purpose of the present study was a comparison of an fish farm (PAL Anlagenbau GmbH, Germany) was stocked in unit II. The
aquaponic cultivation of Nile tilapia red strain (Oreochromis niloticus) 13.11.2013, both fish species were measured, with an initial body
and African catfish (Clarias gariepinus) on plant growth. Two identical weight of 3.32 g (±1.63) for O. niloticus (n = 33) and 3.92 g (±1.63)
 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63 53

Fig. 1. Schematic overview of the two identical built coupled aquaponic units at University of Rostock (F = fish tank; C = clarifier; Pb I–V = plant boxes 1–5 with drip overhead irrigation
system and a bell siphon outlet (●) at the end of each box and 9 PVC test tubes (Pb I, III, V, ○) at front, middle and back position, n = 3; S = sump; P = pump).

for C. gariepinus (n = 33, Table 1), and an initial biomass for Nile tilapia
of 2917.3 g (±21.6) and 2934.7 g (±10.5) for African catfish.
Fish were constantly fed with Tilapia feed Aller Aqua 37/10 float
(crude protein: 37%, crude fat: 10%, NFE: 36.7%, ash: 6.9% and fibre:
4.4%, Emsland-Aller Aqua GmbH, Germany) with a particle size of
3 mm over 24 h, using automatic belt feeders (Aquacultur Fischtechnik
GmbH, Germany). The daily feed input was 70 g for both fish species
during the entire run-in phase according to Palm et al. (2014a), and in-
creased to 100 g, with a further increase of 50 g per week until 450 g d−1
at the end of the experiment (Fig. 4). According to physical water pa-
rameters, the experiment contained three phases. During the run-in
phase (days 1 to 14) fish were fed with a total feed input of 1010.00 g
(7.14 % feed d−1) or 2.47% feed per body weight d− 1 for O. niloticus
(2.46% feed C. gariepinus), during the exponential phase (days 15 to
27) with 1650.00 g (7.69% feed d− 1) or 4.35% feed body weight d− 1
for O. niloticus (4.32 % feed C. gariepinus) and during the accumulation
phase (days 28 to 70) with 13,650.00 g (2.33% feed d−1) or 10.88%
feed body weight d−1 for O. niloticus (10.82% feed C. gariepinus). Total
feed amount in both aquaponic systems was 16,310.00 g with an aver-
age daily feed input of 1.45% (236.38 g). For growth evaluation, fish
were weighed every 14 days (n = 33).
Three weeks before the experiment, three different plant species
(basil-Ocimum basilicum [Kiepenkerl, Baldur Garten GmbH, Germany],
parsley-Petroselinum crispum [Mooskrause 2/Smaragd, Sperli GmbH,
Everswinkel, Germany] and marjoram-Origanum majorana [Kiepenkerl,
Nebelung GmbH, Germany]) were seeded in rockwool cubes. The
13.11.2013, the herbs were planted into the hydroponics, with n = 12
per species in each box (n = 36 plants per box, N = 180 plants per
unit) at a distance of 25 cm. For plant growth, no fertilisers or pesticides
were used.

Fig. 2. Aquaponic system at University of Rostock, A: herbs in the gravel hydroponic subsystem (plant box) with PVC test tube and the bell siphon outlet, B: overhead drip irrigation system
with PVC test tubes and the bell siphon, C: herbs from unit I box III, D: herbs from unit II box I after day 70.
54 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63

Table 1 14 days (n = 33). The specific growth rate (SGR% d−1) was calculated
Fish growing parameters and plant final fresh weight over 70 consecutive days (means, with:
±SD), different letters in the same row showing different groups (p b 0.05).

Parameters O. niloticus (unit I) C. gariepinus (unit II) SGR (% d−1) = (ln Wt − ln W0) × t−1 × 100,
Initial body weight (g) 3.32a ± 1.63 3.92a ± 1.63 with Wt = final biomass, W0 = initial biomass, t = time in days.
Final body weight (g) 27.60a ± 21.12 46.29b ± 26.24 The feed conversion ratio (FCR) was calculated with:
Initial body length (cm) 5.8a ± 1.1 8.0b ± 1.2 FCR = fish feed quantity (g) × weight gain (g)−1.
Final body length (cm) 10.6a ± 2.6 17.6b ± 3.6
Initial biomass (g) 2917.3a ± 21.6 2,934.7a ± 10.5
Final biomass (g) 20,966.7a ± 1583.0 29,573.7b ± 453.7 Differences of fish growth, water parameters in the fish tank, the clar-
Initial stocking density (kg m−3) 1.6a ± 0.0 1.6a ± 0.0 ifier and the sump are given in means of aquaponic units I and II. If the
Final stocking density (kg m−3) 11.7a ± 0.9 16.4b ± 0.3 Shapiro-Wilk test showed normality, independent t-test was used; other-
Biomass weight gain (g) 18,049.3a ± 1595.7 26,639.0b ± 454.5 wise the Mann-and-Whitney test identified differences (p b 0.05). Values
SGR (% d−1) 2.81a ± 0.12 3.30b ± 0.02
FCR 0.91a ± 0.08 0.61b ± 0.01
inside the hydroponic plant boxes were analysed by using One Way
Mortality (%) 1.60 10.42 Analyses Of Variance (ANOVA, p b 0.05) and post hoc Tukey-HSD test
Cannibalism (%) 0.00 9.62 (variance homogeneity) or Dunnett-T3 test (variance inhomogeneity).
All data were analysed by using Microsoft Excel (Microsoft®, 2010) and
Basil fresh weight (g) 0.30a ± 0.37 0.12b ± 0.21
SPSS 20.0 (IBM, 2011) statistical software package.
Parsley fresh weight (g) 0.55a ± 0.39 0.27b ± 0.31
Marjoram fresh weight (g) 0.01a ± 0.02 0.00a ± 0.01
3. Results

2.3. Physical and chemical water parameters 3.1. Fish and plant growth

Physical water parameters were taken daily from days 7 to 70 from
the fish tank, the clarifier and the sump of each aquaponic unit at the
same position. Hydroponic plant boxes of each aquaponic unit were
measured on four days during the experiment, with day I = day 21
(03.12.2013), day II = day 35 (17.12.2013), day III = day 49
(31.12.2013) and day IV = day 63 (14.01.2014). Data were collected
from the PVC test tubes of plant boxes I, III and V. Physical parameters
of temperature (°C), dissolved oxygen (DO, mg L−1), oxygen saturation
(%), conductivity (μs cm−1), pH and redox potential (mv) were mea-
sured with a HQ40D multimeter (Hach Lange GmbH, Germany). Chem-
ical water parameters of NH3-N (mg L−1), NO2-N (mg L−1), NO3-N (mg
L−1) and phosphorus (mg L−1) were analysed twice a week with a
spectral photometer DR-3900 (Hach Lange GmbH, Germany). Light
was measured one day per week at the same time (12:00) above the
PVC test tubes of the hydroponic plant boxes (n = 27) with a Digital
Lux Meter (Dr. Meter DM-LX1330B). Photosynthetically Active Radia-
tion light (PAR, μm m−2s− 1) was taken by a Solar Electric Quantum
Meter (Spectrum Technologies, Inc., USA).
No significant differences were found in initial fish growing param-
eters (initial body weight, initial biomass, and initial stocking density,
Table 1). Final growth performance was significantly better in C.
gariepinus (unit II, Fig. 3) with final body weight, final biomass, final
stocking density and biomass weight gain (46.29 g ± 26.24; 29,573.7
g ± 453.7; 16.4 kg m−3 ± 0.3; 26,639.0 g ± 454.5) in contrast to O.
niloticus (27.60 g ± 21.12; 20,966.7 g ± 1583.0; 11.7 kg m−3 ± 0.9;
18,049.3 g ± 1595.7, Table 1). The specific growth rate (SGR) was signif-
icantly better for C. gariepinus with 3.30% d−1 (±0.02) compared with
O. niloticus with 2.81% d− 1 (± 0.12). The feed conversion ratio was
good for both fish species with significant better values for C. gariepinus
(0.61 ± 0.01) in contrast to O. niloticus (0.91 ± 0.08). Survival was high
in both species with a mortality of 1.60% in O. niloticus and 10.42% in C.
gariepinus. Only the African catfish showed cannibalism (9.62%).
Plant growth of herbs was generally low but significantly better in O.
niloticus (unit I) with 0.30 g (±0.37) in basil and 0.55 g (±0.39) in parsley
compared with C. gariepinus (0.12 g ± 0.21; 0.27 g ± 0.31). Marjoram
performed very poor in both units with 0.01 g (±0.02) in unit I with O.
niloticus and 0.00 g (±0.01) in unit II with C. gariepinus (Table 1).

2.4. Statistical analyses 3.2. Chemo-physical water parameter

The dimensionless aquaponic growth factor (AGF) is newly intro- Water parameters were significantly different in dissolved oxygen
duced, representing the deviation from the best possible growth (DO), oxygen saturation, light intensity and Photosynthetically Active
(DFIGF), calculated as a quotient (with DFIGF ideal = 1.00 if final bio- Radiation light (PAR) between the two aquaponic units (Table 2).
mass of fish and plants [g] were equal) of the maximum achieved fish
and plant yield (biomass in g) during an experiment, under comparison
of two identical built aquaponic units or experimental conditions. The
AGF is described as the absolute value of a number (|x|) and compares
the biomass development directly. For calculation of the AGF, the fol-
lowing formula was used:

AGFfish factor I = |((fish I final biomass of unit I) × (fish II final biomass

of unit II)−1) − DFIGF|,
AGFfish factor II = |((fish II final biomass of unit II) × (fish I final bio-
mass of unit I)−1) − DFIGF|,
AGFplant factor I = |((plant biomass of unit I) × (plant biomass of unit
II)−1) − DFIGF|,
AGFplant factor II = |((plant biomass of unit II) × (plant biomass of
unit I)−1) − DFIGF|,
with DFIGF = 1.00.

Fish growth parameters were calculated with initial and final fish Fig. 3. Weight gain of ungraded O. niloticus (aquaponic unit I) and C. gariepinus (aquaponic
biomasses (n = 3) and individual fish body weight and length every unit II) over 70 consecutive days.
 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63 55

Table 2 Table 3
Global chemo-physical water parameters (means, ±SD), different letters in the same row Global physical water parameters in phases I, II and III (means, ±SD), different letters in
showing different groups (p b 0.05). the same row showing different groups (p b 0.05).

Parameters O. niloticus (unit I) C. gariepinus (unit II) Parameters O. niloticus (unit I) C. gariepinus (unit II)

Dissolved oxygen (mg L−1) 7.0a ± 0.9 7.5b ± 0.6 Phase I (day 7 until 14)
Oxygen saturation (%) 81.9a ± 9.4 87.1b ± 5.9 Dissolved oxygen (mg L−1) 7.7a ± 0.3 7.9a ± 0.3
Temperature (°C) 23.1a ± 1.3 22.8a ± 1.3 Oxygen saturation (%) 91.5a ± 1.9 93.3a ± 1.0
pH 7.7a ± 0.4 7.8a ± 0.4 Temperature (°C) 23.7a ± 0.9 23.4a ± 0.9
Electric conductivity (μs cm−1) 884.3a ± 105.9 868.1a ± 107.6 pH 8.2a ± 0.1 8.2a ± 0.1
Redox potential (mv) 208.3a ± 25.3 208.6a ± 23.7 Electric conductivity (μs cm−1) 898.4a ± 19.6 891.9a ± 18.2
Light intensity (lx) 2,259.9a ± 995.1 2,025.8b ± 941.5 Redox potential (mv) 241.2a ± 10.1 241.1a ± 10.8
PAR (μmol m−²s−1) 42.1a ± 17.3 36.9b ± 16.2
NH3-N (mg L−1) 0.24a ± 0.11 0.18b ± 0.07 Phase II (day 15 until 27)
Nitrite (mg L−1) 0.06a ± 0.05 0.05a ± 0.04 Dissolved oxygen (mg L−1) 8.0a ± 0.2 8.1a ± 0.2
Nitrate (mg L−1) 30.94a ± 16.60 23.30a ± 15.16 Oxygen saturation (%) 92.3a ± 2.1 92.3a ± 2.2
Phosphate (mg L−1) 16.86a ± 8.39 11.00b ± 10.10 Temperature (°C) 22.5a ± 1.3 22.1a ± 1.3
pH 8.1a ± 0.1 8.1a ± 0.1
Electric conductivity (μs cm−1) 855.8a ± 124.8 849.8a ± 123.5
Oxygen showed higher levels in aquaponic unit II (C. gariepinus) with Redox potential (mv) 203.8a ± 16.3 206.2a ± 15.8
7.5 mg L− 1 (± 0.6) and 87.1% (± 5.9) in contrast to 7.0 mg L− 1 Phase III (day 28 until 70)
(±0.9) and 81.9% (±9.4) in aquaponic unit I (O. niloticus). Light values Dissolved oxygen (mg L−1) 6.6a ± 0.7 7.2b ± 0.6
were slightly higher in aquaponic unit I (O. niloticus) with a difference of Oxygen saturation (%) 76.8a ± 7.3 84.3b ± 5.1
234.1 lx and 5.2 μmol m−2 s−1 to aquaponic unit II (C. gariepinus). There Temperature (°C) 23.2a ± 1.3 22.9a ± 1.3
pH 7.5a ± 0.4 7.6a ± 0.3
were no significant differences in water temperature at 23 °C in both
Electric conductivity (μs cm−1) 890.0a ± 111.0 868.8a ± 114.4
units; pH was alkaline and between 7.7 and 7.8. Electric conductivity Redox potential (mv) 205.2a ± 26.1 205.0a ± 23.9
(EC) showed values between 884.3 μs cm− 1 in unit I and 868.1 μs
cm− 1 in unit II, the redox potential was comparable at 208 mv.
R² = 0.14; unit II: y = − 0.0269x + 8.6406, R² = 0.25) and in phase
Chemical water parameters were significantly different in ammonia
III decreasing (unit I: y = − 0.0476x + 8.8349, R² = 0.67; unit II:
with 0.24 mg L− 1 (± 0.11) NH3-N in unit I with O. niloticus and 0.18
y = −0.0327x + 8.784, R² = 0.55; Fig. 4).
mg L− 1 (± 0.07) in unit II with C. gariepinus. In contrast, nitrite
was not significant different with 0.06 mg L− 1 (± 0.05) in unit I and
0.05 mg L−1 (± 0.04) in unit II as well as nitrate with 30.94 mg L− 1 3.3. Water parameter in the hydroponic units
(± 16.60) in unit I and 23.30 mg L−1 (± 15.16) in unit II. Phosphate
was significantly different with 16.86 mg L− 1 (± 8.39) in unit I and The water parameter of the hydroponic units reflected in general the
11.00 mg L−1 (±10.10) in unit II. trends for the whole aquaponic units on the four sampling days. De-
During the duration of the experiment, oxygen values were moni- creasing linear trends were recorded for dissolved oxygen, oxygen sat-
tored and statistically analysed in detail, distinguishing the three phases uration and alkaline pH (Table 4; Fig. 5). In contrast, values for
according to an increasing, stagnating and decreasing oxygen level. phosphate showed significant increasing trends between the four days
Phase I was identified until day 14, phase II from day 15 to day 27 and in unit I from 6.0 mg L− 1 (± 0.7) on day I to 21.9 mg L−1 (± 2.9) on
phase III from day 28 to day 70, respectively (Fig. 4). Comparing both day IV and in unit II from 3.1 mg L− 1 (± 0.2) on day I to 8.9 mg L− 1
units, there were no significant differences between all water parame- (±2.2) on day IV. Temperature inside the plant boxes was more stable
ters in phase I and II (Table 3), whereas phase III showed significant dif- in unit I between 23.1 °C and 24.0 °C and significantly lower in unit II
ferences only in dissolved oxygen and oxygen saturation levels. In phase from 21.8 °C to 22.9 °C. Electric conductivity (EC) showed significant dif-
I, the linear equation of dissolved oxygen was increasing (unit I: y = ferences on all days in each of the aquaponic units with a maximum in
0.1154x + 6.5491, R² = 0.91; unit II: y = 0.0957x + 6.9436, R² = unit I on day IV of 1027.7 μs cm−1 and a minimum on day II of
0.91), in phase II slightly decreasing (unit I: y = −0.0206x + 8.4414, 776.3 μs cm−1 (Table 4). Maximum conductivity was observed in unit

Fig. 4. Oxygen distribution (mg L−1) and feed input (g d−1) divided in phase I (run in phase), II (exponential phase), III (accumulation phase), taken from day 7.
56 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63

Table 4
Physico-chemical parameters inside the hydroponic unit (pvc test tubes, means, ±SD n = 27) of unit I (O. niloticus) and unit II (C. gariepinus) on days I, II, III and IV, different letters in the
same row (a, b, c, d) showing significant differences (ANOVA, p b 0.05) between values of one independent unit (time series, along all days); different letters in the same column (m, z)
showing significant differences (t-test, p b 0.05) in comparison of units I and II on the specific experimental day and parameter.

Parameters Day I (21) Day II (35) Day III (49) Day IV (63)

O. niloticus (unit I)
Oxygen (mg L−1) 7.6a, m ± 0.3 6.5b, m ± 0.4 5.5c, m ± 0.4 4.6d, m ± 0.6
Oxygen saturation (%) 88.1a, m ± 3.9 76.8b, m ± 4.2 64.0c, m ± 4.8 61.3d, m ± 3.9
Temperature (°C) 23.1a, m ± 1.1 24.0b, m ± 0.4 23.1a, m ± 0.3 23.9b, m ± 0.5
pH 8.0a, m ± 0.1 7.7b, m ± 0.1 7.4c, m ± 0.0 7.0d, m ± 0.0
Electric conductivity (μs cm−1) 984.6a, m ± 11.1 776.3b, m ± 22.6 895.5c, m ± 13.4 1,027.7d, m ± 8.2
Phosphorus (mg L−1) 6.0a, m ± 0.7 9.7b, m ± 1.1 16.3c, m ± 3.4 21.9d, m ± 2.9

C. gariepinus (unit II)

Oxygen (mg L−1) 7.4a, z ± 0.4 6.7b, z ± 0.5 5.9c, z ± 0.8 5.0d, z ± 0.6
Oxygen saturation (%) 83.6a, z ± 8.6 77.7b, z ± 6.1 67.0c, z ± 9.8 59.3d, m ± 9.2
Temperature (°C) 22.2a, z ± 0.8 22.9b, z ± 0.8 21.8c, z ± 0.9 22.5a, z ± 0.9
pH 8.0a, m ± 0.1 7.8b, z ± 0.1 7.5c, z ± 0.1 7.3d, z ± 0.1
Electric conductivity (μs cm−1) 982.0a, z ± 6.3 769.5b, z ± 5.0 900.2c, z ± 6.2 964.9d, z ± 4.5
Phosphorus (mg L−1) 3.1a, z ± 0.2 4.4b, z ± 1.1 7.9c, z ± 1.5 8.9d, z ± 2.2

II on day I with 982.0 μs cm−1 and minimum with 769.5 μs cm−1 on day
II. No significant differences were only found in pH on day I with 8.0
(±0.1) in unit I and 8.0 (±0.1) in unit II, as well as oxygen saturation
on day IV with 61.3% (± 3.9) in unit I and 59.3% (± 9.2) in unit II
(Table 4).
The plant boxes (I, III, V) of each separate aquaponic unit with the
different PVC test tube positions (front, middle, back; n = 3, Tables 5,
6, 7, 8) were tested for water parameter significance on the four days
of data recording. No clear trend of the recorded water parameters
were found in the plant boxes of hydroponic unit I (O. niloticus). Plant
box I showed significant differences (p b 0.05) of dissolved oxygen
(DO) and oxygen saturation between the front, middle and back posi-
tions only on day I (Table 5). On day IV DO were significant in plant
box V between the test tube positions (Table 6). Temperature showed
significant differences between the positions in aquaponic unit I in
plant box I on day II (Table 5), in plant box V on day I (Table 5) and in
plant box I on day III (Table 6).
Similar, plant boxes of aquaponic unit II with C. gariepinus showed
no clear trend of water parameters. Dissolved oxygen showed signifi-
cant differences in plant box V on days I and II (Table 7). Temperature
was significant between the positions of all plant boxes on day I and
on day II in plant boxes I and V (Table 7). On day IV, significant differ-
ences in temperature were found only in plant box III and V (Table 8).
Levels of pH were significant between the tube positions in plant box I
on day II (Table 7). Conductivity showed significant differences in
plant box III on day II (Table 7).
Fig. 5. Oxygen distribution (mg L−1) of the hydroponic units on four experimental days (I,
II, III, IV). Significant differences are shown in groups of one unit between the days with a,
b, c, d (ANOVA, p b 0.05), and between unit I and unit II on one experimental day with m
and z (t-test, p b 0.05).
4. Discussion
The present study describes I) the performance of coupled small-
scale sized aquaponic units and II) effects of Nile tilapia (O. niloticus)
and African catfish (C. gariepinus) on growth of different herbs under
winter conditions in the northern hemisphere. The units performed sta-
ble during the run of the experiment under continuously increasing
feed input levels from 70 g–450 g per day. As earlier described by
Palm et al. (2014a), and by using oxygen levels as an indicator for sys-
tem stability, we could distinguish the run-in phase (days 1–14) with
low nutrient loads, fish and plant weights (Fig. 4). This phase showed
increasing values of oxygen, phosphate and conductivity. After reaching
a maximum oxygen level, the exponential phase (days 15–27) showed
increasing feed input levels, fish and plant weights, until a distinct drop
in oxygen levels occurred. The conductivity, salinity and nutrient loads
also increased during this phase. Finally, during the following accumu-
lation phase (days 28–70), the increasing feed input, fish and plant bio-
mass resulted in a continuous decreasing oxygen level under increasing
conductivity. The steady phase according to Palm et al. (2014a), where
the feed input balanced the oxygen level, salinity, conductivity and nu-
trient load, was short in the present study and estimated at the feed
input level of 150 g (days 21–27). Fish growth of O. niloticus and C.
gariepinus was different with varying yields of associated herbs.
4.1. Fish growth
Fish growing parameters demonstrated good results for both species
and showed no significant differences in initial body weights, initial bio-
mass and initial stocking density for the Nile tilapia (O. niloticus) and the
African catfish (C. gariepinus, Table 1). Growth of O. niloticus was compa-
rable to Palm et al. (2014a) with a similar experimental aquaponical de-
sign. The daily fish feed input increased over a period of 160 days,
younger O. niloticus (0.50 g initial weight) showed comparable specific
growth rates (2.98–3.04% d−1) and feed conversion between 0.83 and
0.93 as found in the present study. With the use of African catfish, fish
growing parameter differed according to the species-specific growth
potential (Fig. 3). Specific growth rate (SGR) and feed conversion ratio
was significant better for C. gariepinus with 3.30% d−1 (±0.02) and 0.61
(±0.01), respectively (Table 1), and better compared with earlier studies
under aquaponic production and stable feed amounts per day. Older C.
gariepinus with an initial body weight of 480.23 g (±75.68) reared in a
similar aquaponic system and a stable feed input of 200 g d−1 showed re-
duced specific growth of 0.65% d−1 with the production of lettuce,
cucumber, tomato and basil (Palm et al., 2014b). Endut et al. (2010) re-
ported lower specific growth rates between 1.68 and 1.83% d−1 in a grav-
el substrate aquaponic system with juvenile C. gariepinus (30–40 g) held
under different hydraulic loading rates, a feeding amount of 2–4% body
 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63 57

Table 5
Physico-chemical parameters within the plant boxes at the specific positions (front, middle, back) of unit I (O. niloticus) on days I and II, means (±SD, n = 3), different letters in groups
showing significant differences (ANOVA, p b 0.05).

Position parameters Box Front Middle Back Front Middle Back

Day 1 Day 2
Oxygen (mg L−1) I 7.7a ± 0.1 7.8b ± 0.0 7.9c ± 0.1 6.3a ± 0.3 6.5a ± 0.1 6.6a ± 0.1
Oxygen (%) 87.9a ± 0.7 90.7b ± 0.4 92.1c ± 0.5 73.3a ± 4.3 77.4ab ± 1.6 78.7b ± 1.6
Temperature (°C) 22.7a ± 0.2 24.2ab ± 3.0 23.3b ± 0.2 23.3a ± 0.2 24.3b ± 0.1 24.5c ± 0.1
pH 8.1a ± 0.0 7.9ab ± 22.0 8.1b ± 0.0 7.7a ± 0.1 7.7a ± 0.2 7.7a ± 0.2
Conductivity (μs cm−1) 982.7a ± 4.2 989.1a ± 4.9 990.7a ± 9.9 775.7a ± 1.7 775.6a ± 1.3 773.8a ± 1.9
Phosphate (mg L−1) 6.0a ± 0.7 6.2a ± 0.5 5.9a ± 0.6 9.1ab ± 1.2 8.5a ± 0.5 9.7b ± 0.6
Oxygen (mg L−1) III 7.4a ± 0.2 7.6a ± 0.2 7.4a ± 0.6 6.3a ± 0.3 7.0b ± 0.2 6.1a ± 0.4
Oxygen (%) 84.9a ± 2.2 88.6b ± 2.9 85.4ab ± 6.3 73.8a ± 3.0 83.6b ± 2.0 72.2a ± 4.5
Temperature (°C) 22.7a ± 0.1 23.4b ± 0.1 22.8a ± 0.2 23.6a ± 0.1 24.2b ± 0.2 24.1b ± 0.1
pH 7.9a ± 0.1 8.1b ± 0.0 8.0ab ± 0.1 7.7a ± 0.2 7.8a ± 0.1 7.8a ± 0.0
Conductivity (μs cm−1) 985.1a ± 3.2 988.4a ± 4.7 976.3a ± 16.2 794.2a ± 67.5 775.3a ± 3.0 769.8a ± 2.3
Phosphate (mg L−1) 5.6a ± 0.8 5.8a ± 0.2 5.9a ± 0.4 9.5a ± 1.3 10.1a ± 1.0 9.1a ± 1.2
Oxygen (mg L−1) V 7.7a ± 0.1 7.9b ± 0.1 7.3c ± 0.3 6.8a ± 0.3 6.6ab ± 0.1 6.4b ± 0.2
Oxygen (%) 88.2a ± 1.5 91.3b ± 1.4 83.4c ± 3.1 78.4a ± 0.8 78.4a ± 1.3 75.4b ± 2.8
Temperature (°C) 22.7a ± 0.2 23.2b ± 0.2 22.9c ± 0.2 23.7a ± 0.2 24.4b ± 0.1 23.9a ± 0.3
pH 8.1a ± 0.0 8.1a ± 0.0 7.8b ± 0.2 7.9a ± 0.0 7.8a ± 0.1 7.6b ± 0.1
Conductivity (μs cm−1) 993.9a ± 12.3 984.7a ± 2.6 970.6b ± 12.1 777.2a ± 3.3 776.9a ± 3.6 768.4b ± 4.4
Phosphate (mg L−1) 6.6a ± 1.3 6.2ab ± 0.3 5.6b ± 0.5 10.9a ± 0.5 10.1b ± 0.2 10.1b ± 0.3

weight d−1 and the cultivation of water spinach (Ipomoea aquatica). Sim-
ilarly, Sikawa & Yakupitiyage (2010) reported a twofold lower specific
growth rate of 1.56% d−1 with the use of nearly two times older African
catfish hybrids (Clarias macrocephalus × Clarias gariepinus) held in an
earthen pond with adapted feed input (10–3% body weight d−1) and
the combined production of lettuce (Lactuca sativa) cultured on three ag-
gregates (gravel, sand, styrofoam).
The feed conversion ratio (FCR), found in the present study, was
more efficient in C. gariepinus in contrast to O. niloticus (Table 1) and de-
scribed in other studies. Under aquaponical production, a much lower
feed conversion ratio of 2.29 was reported by Sikawa & Yakupitiyage
(2010) with only little older C. gariepinus fingerlings (6.58 g ± 1.72)
stocked in an earthen pond. However, juvenile African catfish with an
initial weight of 34.0–118.0 g showed lower feed conversion of 1.23–
1.39 in combination with the production of water spinach (Endut
et al., 2009). In contrast, the feed conversion of much older preadult Af-
rican catfish was more comparable with 1.00 (initial weight: 480.23 g ±
75.68, final weight: 873.77 g ± 165.00, see Palm et al., 2014b). Labora-
tory aquaculture investigations showed a comparable feed conversion
in 4.5 fold younger C. gariepinus (Clarias lazera) with 0.60 (initial weight
0.87 g, final weight 4.33 g) by Hogendoorn (1981) or little lower with
0.73 in two times older fish (initial weight 8.53 g) reported by
Aderolu et al. (2010), and lower with 0.8 and 0.9 (initial weight 82.4–
192.6 g) in much older fish held in tanks or aquaria described by
Martins et al. (2005). Protein content of the feed (37%) has not influ-
enced growth performance of C. gariepinus negatively. According to Ali
& Jauncey (2004), a protein content between 35 and 40% is recommend-
ed for African catfish juveniles (13.45 g ± 0.05) and C. gariepinus accepts
up to 5% feed of their body weight daily. The very good feed conversion
of C. gariepinus in the present study can be explained with the overfeed-
ing condition (up to 10.82% body weight per day) between days 28 and
70 (phase III) based on the initial fish biomass, and caused by the exper-
imental design (increasing feed amount). The fish feeding was very ef-
fective at this time with a rapid growth curve expression according to
the specific fish growth potential, explaining our very good FCR. Further,
extensive stocking density could affect growth of C. gariepinus positive-
ly. After Hossain et al. (1998), best growth of C. gariepinus fry (0.79 g)
was found with 5 fish L−1 under reduced light regime with the use of
cover and shelter. Stocking density in the present study with 0.35 fish
L− 1 (1 fish per 2.89 L−1, 346 fish m−3, 624 fish 1.8 m−3) was lower
but mortality was comparable to 11.15% (10.42%, Table 1). With the
very low stocking density and natural light regime in our study light
shading effects could be compensated and future investigations are
needed to point out this effects under aquaponic production conditions.

Table 6
Physico-chemical parameters within the plant boxes at the specific positions (front, middle, back) of unit I (O. niloticus) on days III and IV, means (±SD, n = 3), different letters in groups
showing significant differences (ANOVA, p b 0.05).

Position parameters Box Front Middle Back Front Middle Back

Day III Day IV

Oxygen (mg L−1) I 5.9a ± 0.2 5.6b ± 0.1 5.5ab ± 0.4 4.5a ± 0.2 4.4a ± 0.2 3.5b ± 0.8
Oxygen (%) 67.7a ± 1.9 65.9a ± 1.3 64.6a ± 4.3 59.2a ± 5.5 62.2a ± 2.1 61.2a ± 3.3
Temperature (°C) 22.6a ± 0.3 23.3b ± 0.1 23.5c ± 0.1 23.2a ± 0.3 24.0b ± 0.3 24.1b ± 0.2
pH 7.4a ± 0.0 7.4b ± 0.0 7.4b ± 0.0 7.0a ± 0.0 7.0a ± 0.0 7.0a ± 0.0
Conductivity (μs cm−1) 886.1a ± 28.6 901.8a ± 5.7 896.9a ± 6.6 1027.6a ± 1.5 1028.7a ± 2.0 1027.6a ± 4.6
Phosphate (mg L−1) 22.5a ± 7.6 15.6a ± 1.2 15.8a ± 0.7 22.1a ± 0.8 23.2a ± 2.0 20.6b ± 1.2
Oxygen (mg L−1) III 4.9a ± 0.3 5.6b ± 0.1 5.4b ± 0.3 4.8a ± 0.2 4.8a ± 0.1 4.5b ± 0.0
Oxygen (%) 55.9a ± 3.0 65.7b ± 1.8 62.6b ± 2.9 60.7a ± 3.9 60.7a ± 4.2 60.5a ± 4.4
Temperature (°C) 22.6a ± 0.1 23.2b ± 0.2 23.1b ± 0.1 24.1a ± 0.1 24.4b ± 0.1 24.1a ± 0.0
pH 7.3a ± 0.0 7.4a ± 0.1 7.3a ± 0.0 7.0a ± 0.0 7.1b ± 0.0 7.0a ± 0.0
Conductivity (μs cm−1) 893.9a ± 3.3 901.1b ± 3.7 896.6ab ± 6.3 1029.1ab ± 2.9 1032.0a ± 1.0 1022.4b ± 8.6
Phosphate (mg L−1) 15.3a ± 0.4 14.4b ± 0.7 15.9a ± 1.3 20.1a ± 1.8 20.8a ± 0.6 20.3a ± 0.8
Oxygen (mg L−1) V 5.8a ± 0.3 5.7a ± 0.3 5.1b ± 0.4 5.0a ± 0.1 4.7b ± 0.2 5.3c ± 0.3
Oxygen (%) 66.8a ± 4.0 66.9a ± 3.2 59.7b ± 4.6 66.1a ± 2.8 59.3b ± 1.5 61.6b ± 2.3
Temperature (°C) 22.9a ± 0.1 23.3b ± 0.1 23.1ab ± 0.2 23.3a ± 0.7 24.0b ± 0.4 23.8ab ± 0.2
pH 7.4a ± 0.0 7.4b ± 0.0 7.4ab ± 0.0 7.0a ± 0.0 7.0a ± 0.0 7.1b ± 0.0
Conductivity (μs cm−1) 898.1ab ± 6.8 902.6a ± 4.2 882.1b ± 18.0 1033.3a ± 10.9 1030.9a ± 2.8 1017.9a ± 14.9
Phosphate (mg L−1) 15.1a ± 1.2 16.0a ± 0.4 16.4a ± 0.8 28.5a ± 2.1 21.0b ± 0.9 20.8b ± 1.8
58 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63

Table 7
Physico-chemical parameters within the plant boxes at the specific positions (front, middle, back) of unit II (C. gariepinus) on days I and II, means (±SD, n = 3), different letters in groups
showing significant differences (ANOVA, p b 0.05).

Position parameters Box Front Middle Back Front Middle Back

Day I Day II
Oxygen (mg L−1) I 7.1a ± 0.1 7.7b ± 0.1 7.5ab ± 0.4 6.3a ± 0.1 7.0b ± 0.2 7.0b ± 0.1
Oxygen (%) 80.1a ± 1.1 89.1b ± 0.9 86.3b ± 4.8 72.1a ± 0.9 81.9b ± 2.5 82.4b ± 0.5
Temperature (°C) 21.9a ± 0.2 23.2b ± 0.1 23.0c ± 0.1 22.6a ± 0.2 23.8b ± 0.1 23.5c ± 0.3
pH 8.0a ± 0.1 7.9a ± 0.1 8.1b ± 0.0 7.8a ± 0.0 7.8b ± 0.0 7.9c ± 0.0
Conductivity (μs cm−1) 985.1a ± 4.5 986.8a ± 4.2 984.8ab ± 2.2 770.6ab ± 8.9 766.00a ± 0.9 769.9b ± 0.9
Phosphate (mg L−1) 3.0a ± 0.1 3.1b ± 0.1 3.1bc ± 0.1 3.7a ± 0.2 3.6a ± 0.2 4.6a ± 1.4
Oxygen (mg L−1) III 7.8a ± 0.0 7.8a ± 0.1 7.5b ± 0.2 7.0a ± 0.2 7.1a ± 0.1 7.1a ± 0.2
Oxygen (%) 87.3a ± 0.3 89.5b ± 1.9 85.6a ± 2.1 80.4a ± 2.2 82.7b ± 1.1 82.3ab ± 2.0
Temperature (°C) 21.9a ± 0.4 22.8b ± 0.2 22.3c ± 0.5 22.7a ± 0.1 23.5b ± 0.2 23.3b ± 0.3
pH 8.1a ± 0.0 8.1a ± 0.0 8.1a ± 0.1 7.9a ± 0.0 7.8a ± 0.3 7.9a ± 0.0
Conductivity (μs cm−1) 979.6a ± 4.9 985.9a ± 7.4 979.2a ± 2.5 763.00a ± 1.1 773.2b ± 4.1 768.7c ± 2.9
Phosphate (mg L−1) 3.1a ± 0.1 3.1a ± 0.1 3.1a ± 0.2 3.9a ± 0.3 4.5a ± 0.5 5.1a ± 2.5
Oxygen (mg L−1) V 6.7a ± 0.1 7.6b ± 0.1 7.0c ± 0.2 5.9a ± 0.1 6.8b ± 0.1 6.2c ± 0.3
Oxygen (%) 69.0a ± 17.3 87.0b ± 0.7 78.4a ± 2.1 67.5a ± 3.2 79.5b ± 1.0 69.9a ± 2.9
Temperature (°C) 21.0a ± 0.2 22.7b ± 0.4 21.3c ± 0.2 21.7a ± 0.1 23.5b ± 0.1 21.9c ± 0.1
pH 8.0a ± 0.0 8.0a ± 0.1 8.0a ± 0.1 7.7a ± 0.0 7.6b ± 0.1 7.8a ± 0.0
Conductivity (μs cm−1) 974.7a ± 1.9 987.2b ± 4.3 974.9a ± 3.8 768.3a ± 2.6 771.1ab ± 4.3 774.7b ± 2.1
Phosphate (mg L−1) 3.2ab ± 0.4 3.4a ± 0.3 2.9b ± 0.3 4.7a ± 0.2 5.1b ± 0.5 4.8ab ± 0.1

For O. niloticus, specific growth was below that observed for C.
gariepinus, and within the normal range described for aquaponics and
under aquaculture production. In standalone aquaculture, Likongwe et
al. (1996) reported slightly reduced growth of 2.24% d−1 under little
higher water temperature of 24 °C and little older fish of 4.75 g, which
is in the normal range of growth. At a much higher temperature of
32 °C in the range of optimal growth, these fish showed a specific
growth similar to the present study and our younger fish with 2.82%
d−1. However, feeding management and feed composition was better
by Likongwe et al. (1996) with 3% feed per body weight (by hand
once per day) and 50% protein in the feed in contrast to the present
study with a feeding interval of 24 h by automatic belt feeders, a
lower average daily feed input of 1.45% and 37% crude protein. Recom-
mended protein proportions were described to be 45% for 0.4–0.5 g fry
and 35% for 17–22 g fingerling and advanced juvenile of 37–43 g
(Abdel-Tawwab et al., 2010) or 40% for 0.51 g fry and 30% for larger
fish of 96–264 g (Al Hafedh, 1999) and were slightly higher as used in
the present experiment with an initial weight of 3.32 g of O. niloticus.
However, specific growth rate of 2.89% d− 1 described by Al Hafedh
(1999) was comparable with a lower protein content of 30% and youn-
ger O. niloticus of 0.51 g but with a feeding rate of 15–3% over 140 days.
We fed O. niloticus with steadily increasing feed input from 2.47 to
10.88% per body weight; a contrary feeding management depended
on the experimental design with resulting effects on specific growth.
Compared to raft aquaponics, lower specific growth rates of 1 and
1.1% d− 1 (mixed sex, all male population) were described by Al-
Hafedh et al. (2008) with different older initial fish weights in staggered
production (initial weights of 42.5 g, 74.8 g, 138 g, 248 g) and the pro-
duction of leaf lettuce. The feed utilisation of O. niloticus in our study
can be explained with primary reduced, later highly increased and in
general restricted feed amounts. In raft aquaponics, Al-Hafedh et al.
(2008) reported little lower FCRs of 1.3–1.4 with staggered all male
and mixed sex O. niloticus population of older fish. Similarly, Rakocy et
al. (2004) found FCRs of 1.7 for the Nile tilapia (79.2 g) and 1.8 for the
Red tilapia (58.8 g) in combination with batch and staggered production
of basil and okra in raft aquaponics. Further, older O. niloticus (initial
weight: 173.51 g ± 14.64, final weight: 337.63 g ± 54.04) held in the
same modified system, also under winter time conditions (T ≈ 23–
24 °C), had a lower FCR of 1.03 (Palm et al., 2014b). In general, feed con-
version of O. niloticus was better than expected but reduced compared
to the FCR of C. gariepinus. The growth curve of O. niloticus at the begin-
ning of the experiment was more reduced with the same feed input for
both fish species and showed only a moderate weight gain until the end
of the study (Fig. 3).
For both fish species, chemo-physical water parameters were in its
optimal range (Table 2). Dissolved oxygen concentrations were

Table 8
Physico-chemical parameters within the plant boxes at the specific positions (front, middle, back) of unit II (C. gariepinus) on days III and IV, means (±SD, n = 3), different letters in groups
showing significant differences (ANOVA, p b 0.05).

Position parameters Box Front Middle Back Front Middle Back

Day III Day IV

Oxygen (mg l−1) I 6.8a ± 0.1 6.4ab ± 0.5 6.4b ± 0.2 4.8a ± 0.2 4.9a ± 0.3 5.3b ± 0.2
Oxygen (%) 77.8a ± 1.0 73.7ab ± 5.7 73.8b ± 2.0 54.3a ± 2.8 61.4b ± 1.6 62.5b ± 3.0
Temperature (°c) 22.2a ± 0.1 22.6b ± 0.1 22.7b ± 0.2 22.2a ± 0.4 23.5b ± 0.1 23.5b ± 0.1
ph 7.5a ± 0.0 7.4b ± 0.0 7.5a ± 0.0 7.3a ± 0.0 7.2b ± 0.0 7.2b ± 0.0
Conductivity (μs cm−1) 899.0a ± 5.9 898.9a ± 2.1 901.0a ± 3.7 964.9a ± 1.3 965.0a ± 0.7 968.3a ± 7.2
Phosphate (mg l−1) 7.3a ± 0.5 6.9a ± 0.4 7.7b ± 0.4 8.2a ± 0.6 9.9b ± 0.9 8.8a ± 0.6
Oxygen (mg l−1) III 6.5a ± 0.1 5.6b ± 0.8 5.9ab ± 1.1 5.3a ± 0.1 5.6b ± 0.1 5.5b ± 0.2
Oxygen (%) 73.8a ± 1.5 64.0b ± 8.1 66.9ab ± 12.3 66.9a ± 3.3 67.6a ± 3.4 70.1a ± 1.3
Temperature (°c) 21.8a ± 0.1 21.9a ± 0.9 22.0a ± 0.4 22.3a ± 0.2 23.2b ± 0.2 22.9c ± 0.2
ph 7.7a ± 0.0 7.6b ± 0.0 7.5b ± 0.0 7.4a ± 0.0 7.3b ± 0.0 7.3b ± 0.1
Conductivity (μs cm−1) 904.6a ± 5.3 908.6a ± 1.4 902.3a ± 7.0 962.8a ± 3.7 966.2b ± 2.0 964.2ab ± 2.1
Phosphate (mg l−1) 9.4a ± 2.6 8.5a ± 1.3 7.5a ± 0.8 9.5a ± 1.6 7.8a ± 0.8 7.8a ± 2.4
Oxygen (mg l−1) V 4.9a ± 0.2 5.7b ± 0.3 4.9a ± 0.3 4.4a ± 0.2 4.6a ± 1.0 4.2a ± 0.5
Oxygen (%) 53.9a ± 2.3 65.3b ± 3.4 54.5a ± 2.6 49.5a ± 2.5 54.2a ± 12.0 47.1a ± 5.3
Temperature (°c) 20.2a ± 0.1 22.4b ± 0.3 20.4a ± 0.2 20.9a ± 0.1 22.9b ± 0.3 21.2c ± 0.2
pH 7.5a ± 0.2 7.4a ± 0.0 7.5a ± 0.0 7.2a ± 0.0 7.2b ± 0.0 7.2b ± 0.0
Conductivity (μs cm−1) 891.8a ± 4.3 899.9b ± 3.3 896.0b ± 2.8 960.8a ± 5.5 965.8a ± 4.9 965.9a ± 5.5
Phosphate (mg l−1) 7.3a ± 0.8 8.6a ± 2.2 8.3a ± 1.2 10.7a ± 2.6 8.7a ± 1.8 9.1a ± 4.5
 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63
acceptable with 7.0 mg L− 1 (± 0.9) for O. niloticus and 7.5 mg L− 1
(±0.6) for C. gariepinus. DeLong et al. (2009) suggested an operating
oxygen level between 5.0 and 7.5 mg L−1 for tilapia, whereas chronical-
ly low oxygen concentrations below 3.5 mg L−1 can negatively affect ti-
lapia growth and feed conversion. For C. gariepinus juveniles, with its
air-breathing physiology, oxygen levels below 3.0 mg L− 1 showed a
change to atmospheric air consumption by the fish (Oellermann,
1996). In our study, the minimum oxygen concentration was found at
6.0 mg L−1 in unit II (C. gariepinus) at the end of phase III on day 70
(Fig. 4) with a difference of 3 mg L−1 to the critical DO value. Water
temperature was moderate in the present study with values at 23 °C
in units I and II (Table 2). Acceptable growth for tilapia was reported
at a higher temperature above 25 °C, optimal growth was suggested at
27–29 °C (DeLong et al., 2009). Temperature preference for C. gariepinus
juveniles (1.1 g ± 0.4) was investigated by Oellermann (1996) with a
congregation of a 28–30 °C zone, much higher than in the present
study. Furthermore, an alkaline pH of 7.7 did not influenced O. niloticus
growth; an optimal range of pH between 6 and 9 was reported by
DeLong et al. (2009). The concentration of Ammonia (NH3-N) with
nearly 0.20 mg L− 1 in unit I and II (Table 2) had no negative impact
onto fish growth. Critical levels were reported at 2 mg L− 1 for O.
niloticus (Rakocy, 1989b) and reduced growth of 0.34 mg L− 1 for C.
gariepinus (Schram et al., 2010). Also, nitrite levels of 0.06 (± 0.05)
and 0.05 (± 0.04) mg L− 1 in unit I and II met the requirements of
both fish species with values below 5 mg L−1 for O. niloticus (Rakocy,
1989b) and 0.6 mg L−1 for C. gariepinus (Roques et al., 2015). Nitrate
levels showed no significant differences with 30.94 mg L−1 (±16.60)
in unit I and 23.30 mg L−1 (±15.16) in unit II with no negatives effects
on fish growth (Table 2). After DeLong et al., (2009), much higher ni-
trate levels of 300–400 mg L−1 can affect O. niloticus growth, whereas
C. gariepinus can tolerate nitrate levels up to 140 mg L−1 (Schram et
al., 2014).
4.2. Plant growth
Plant growth was general under expectation in both aquaponic sys-
tems, resulting from environmental conditions during winter. However,
best results were observed for basil (Ocimum basilicum) and parsley
(Petroselinum crispum) in combination with O. niloticus (Table 1). Mar-
joram (Origanum majorana) performed very poor in both aquaponic
systems as a consequence of adaptation problems with the water hy-
droponic environment, light or low temperature. Plant growth in com-
bination with C. gariepinus showed a 2–2.5 fold reduction compared to
the cultivation with O. niloticus. For photoautotrophic organisms light
is the primary growth factor, followed by temperature and nutrient
supply.
Light intensity levels in the present study were in general low. In
aquaponic unit I (O. niloticus), the light intensity was significantly higher
but with small differences of natural illumination (234.1 lx) and Photo-
synthetically Active Radiation light (PAR, 5.2 μmol m−2 s−1), respectively
(Table 2). For greenhouse plant production of the northern latitudes,
Gruda (2005) referred low light intensity as the most limiting climatic
factor. Other environmental factors cannot be used efficiently by plants
if the level of Photosynthetic Photon Flux Density (PPFD) is low during
winter. Glasshouse structure may reduce the daylight quantity to 30% or
more. Short and cloudy days, as they were observed in the present
study, with suboptimal integrals of solar radiation can affect plant growth
and fruit production negatively. This agrees with two experiments by
Succop & Newman (2004) with the cultivation of sweet basil (Ocimum
basilicum) in different cultivation periods. Spring to summer cultivation
(March to July) showed increasing weekly harvests in contrast to basil
production from summer to autumn (August to November) with later de-
clining growth rates in correspondence with decreasing natural light in-
tensities. Under aquaponical production and different electrical night
illumination, Palm et al. (2014b) found no significant differences in basil
gross biomasses of 117.94 g for C. gariepinus (23.33 μmol m² s−1 ±
59
42.32) and 159.00 g for O. niloticus (50.17 μmol m² s−1 ± 49.62). The
study showed no direct growth dependence of light by basil with near
two fold higher PAR level of artificial light at night-time in combination
with the production of O. niloticus. In comparison, PAR climate in the pres-
ent study was slightly higher in aquaponic unit I with 42.1 μmol m² s−1
for O. niloticus, and slightly lower in aquaponic unit II with 36.9 μmol m²
s−1 for C. gariepinus. The difference of PAR between the two aquaponic
units in our study was only 5.2 μmol m² s−1 in comparison to the differ-
ence of 26.84 μmol m² s−1 reported by Palm et al. (2014b). We suggest
that the variation of natural light illumination between the two aquaponic
units had a minor influence onto plant yield, less than the reduced total
natural light levels in wintertime, and light differences alone cannot ex-
plain the better yield of herbs in combination with the production of O.
niloticus. It must be keep in mind that the windows (roof and lateral) of
the glasshouse were limed during summer and autumn to protect plants
against stronger solar irradiation, causing reduced light intensities in
winter.
Water temperature in the present study was close to 23 °C. Rakocy
(2012) suggested an optimal water temperature between 22 and
24 °C for the production of crops in aquaponics. This is in agreement
to temperature optima under aquaponic production for O. basilicum
and P. crispum of 20–25 °C and 15–25 °C (Somerville et al., 2014), re-
spectively. In greenhouse production, O. majorana was grown at tem-
peratures of 21–22 °C (Circella et al., 1995; Banchio et al., 2008). In
aquaponics, higher yields of basil were reported by Palm et al.
(2014b) with a similar water temperature. Poor growth of basil was ob-
served by Kotzen & Appelbaum (2010) at lower temperatures in fresh-
water (21.0–22.1 °C) and brackish water (19.3–20.8 °C). Under
greenhouse production, Succop & Newman (2004) reported generally
better yields of sweet basil (Ocimum basilicum) at a higher temperature
(28.9 °C) and in contrast a higher yield of basil grown in rockwool at a
lower temperature (20.0 °C). Season consequently has a direct influence
on basil growth. Cabanillas et al. (2013) found in Argentina a better in-
season production (October) with the full growth potential of basil in
contrast to the out of season production. In general, basil (O. basilicum)
showed no clear dependence on temperature (low or high) either
grown under standalone hydroponical production nor in aquaponics
but with slightly better growth in freshwater, compared to brackish
water aquaponics with a higher conductivity and slightly reduced tem-
perature. Apparently, minimum temperature level for basil seems to be
20 °C but higher temperature is recommended. This contrasts the
growth of parsley (Petroselinum sp.) that was better under reduced tem-
perature regimes at 20 °C. We can conclude that the temperature in the
present study was in an optimal range for the growth of culinary herbs.
Nutrient dynamics was mainly affected by the continuously increas-
ing fish feed input in combination with different fish species selection
and the retention of nutrients inside the substrate and the plants. The
chemical water parameters met the requirements for culinary herbs
(Table 2). However, better growth was observed inside the aquaponics
with O. niloticus with significant higher levels of NH3-N and phosphate
(P). Nitrite levels in both aquaponic systems were low and nitrate
values much higher in comparison to nitrite. This is an indication for
good oxidative conditions where the oxidation of nitrite to nitrate can
be ten times faster in contrast to the oxidation of ammonia to nitrite
(Van Rijn, 1996), resulting in a higher level of ammonia, a low level of
nitrite and increasing levels of nitrate that is preferentially uptaken by
the plants. Under low nitrate concentration, plants are also able to ad-
sorb nitrogen in the form of ammonium (Weiler & Nover, 2008). With
an increasing feed input similar to the present study, Palm et al.
(2014a) recorded higher levels of TAN (1.12 mg L− 1 unit I, 0.84 mg
L−1 unit II) and lower nitrate values (9.25 mg L−1 unit I, 7.28 mg L−1
unit II) under production of younger O. niloticus and a variety of differ-
ent plants such as herbs and fruiting crops. Comparable levels of nitrate
(31.06 mg L−1) were reported by Palm et al. (2014b) under cultivation
of older O. niloticus in combination with lettuce, cucumber, tomato and
basil, and a stable feed input of 200 g d−1. The fish feed input in the
60 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63
present study was higher and the nutrient retention of the selected
herbs was moderate (see above), resulting in an accumulation of nitrate
and phosphorus. However, the amounts of phosphorus met the demand
for the cultivated plant species as reported by Goddek et al. (2015) for
basil with 8 mg L−1, lower than in aquaponic unit II with C. gariepinus
(11 mg L−1). Lower levels of phosphorus were reported by Palm et al.
(2014b) with a stable feed input of 200 g d− 1, Lennard & Leonard
(2004) with a constant feed input of 215.0 g per day and Lennard &
Leonard (2006) with a feed input of 220.0 g in all systems comparable
to the present study (mean 236.38 g per day).
Physical water parameters, pH, electric conductivity (EC) and redox-
potential were in an optimal range. The pH was more alkaline (7.7–7.8),
optimal for nitrification (Tables 5, 6, 7, 8). The availability of trace metals
is optimal below a pH of 7.0 and for phosphorus, calcium, magnesium,
molybdenum above 6.0 (Rakocy, 2012). As a compromise, Rakocy
(2012) suggested a pH close to 7.0, close to the present study at the
end of the experiment in phase III (Tables 3, 4) and on data sampling
day IV (day 63). Electric conductivity levels between 849 and 898 μs
cm− 1 (Table 3) were little higher as recommended by Rakocy et al.
(2006) for aquaponic systems with 300–600 μs cm−1 (0.3–0.6 mmho
cm−1). Phytotoxic levels in aquaponics are reached at much higher EC
levels above 3500 μs cm−1 (3.5 mmho cm−1, Rakocy et al., 2006).
Other aquaponic systems with good plant growth had higher EC levels.
Palm et al. (2014a) reported no significant EC values of 1018.79 and
975.24 μs cm−1 under cultivation of O. niloticus and different plants.
Higher EC levels near 1200 μs cm−1 were described by Palm et al.
(2014b) without negative effects onto plant growth and the production
of C. gariepinus and O. niloticus. Much higher EC levels were reported by
Kotzen & Appelbaum (2010) in brackish water (about 4500 μs cm−1)
with successful growth of aubergine, celery, chard, chili pepper, chives
(gravel), kohlrabi, parsley (little, small) and mint. In our study, the
redox potential showed no significant differences with values of
208 mv between the two identical aquaponic units (Table 2), a conse-
quence of good water quality. The process of oxidation correlates with
the emission of electrons and the reduction with the admission of elec-
trons mostly found in organic compounds like uneaten feed and excre-
ments. In natural habitats, with a stable relation of oxidation and
reduction, redox potentials of 200 mv occur, similar to the present
study. With higher oxygen levels, the redox potential can reach
300 mv (Sander, 1998). Slightly reduced redox levels of 190 mv were
reported by Palm et al. (2014a) with a comparable increasing feed
input over an experimental time of 160 days in a similar aquaponic sys-
tem and the production of O. niloticus and different plants.
Plant growth was positively affected by the parallel arrangement of
the plant boxes and the hydroponic overhead drip irrigation system
(Figs. 1, 2B) with a decentred nutrient water inflow. No differences
were found between the growths of basil, parsley and marjoram be-
tween the hydroponics of each aquaponic unit. Only basil growth was
reduced in plant box II of aquaponic unit II with the production of C.
gariepinus. Analysed water parameters inside the plant boxes showed
no general trend from the front to the middle and back position,
reflecting the equal water parameter conditions (Tables 5, 6, 7, 8).
This agrees with findings of Palm et al. (2015), who reported similar
conditions in each plant box as a consequence of the parallel arrange-
ment. However, Palm et al. (2015) found a better growth of plants
close to the central water inflow into the hydroponics. In the present
study, the growth of plants was uniform in each box, positively affected
by the decentralised overhead irrigation system. The nutrient enriched
water spread over the entire plant box area, providing an optimal nutri-
ent supply. Dissolved oxygen levels never reached a critical value of 3
mg L−1 as reported by Gislerød & Kempton (1983) for cucumber
(Cucumis sativus) in NFT hydroponics. The lowest oxygen value found
was 3.5 mg L−1 in aquaponic unit I (O. niloticus) and day IV in box I at
the back position. Oxygen levels were lowest inside the hydroponic
units and can be used for monitoring system stability as earlier de-
scribed by Palm et al. (2015).
4.3. Indication of system stability
The concentration of oxygen between unit I (O. niloticus) and unit II
(C. gariepinus) in phase III was significant higher in the C. gariepinus sys-
tem (7.2 mg L−1 = 100%) with a difference of 0.6 mg L−1 (8.34%) to O.
niloticus (6.6 mg L−1 = 91.66%, Table 3). This DO difference could be a
result of the lower oxygen demand of the air breathing African catfish
(C. gariepinus), in accordance with Palm et al. (2014b) who recorded a
2.22 mg L− 1 (32%) difference in oxygen consumption between both
species. While we used smaller sized African catfish (3.92 g), and DO
differences became relevant at the beginning of the accumulation
phase III (day 28) when C. gariepinus had a calculated body weight
and length of approx. 12.9 g and 11.8 cm, Palm et al. (2014b) used
480 g fish. Moussa (1957) already demonstrated that juvenile C.
gariepinus (12.5–32.5 cm) in well-oxygenated water tried to swallow
also atmospheric air. The prevention of free surface access results in as-
phyxia and fish died without an influence of oxygen contents of the
water. Longer survival was recorded for shorter (12.5 cm) compared
with longer fish (32.5 cm). In contrast, Oellermann (1996) observed
from existing air breathes at 6 mg L−1 an increasing air breathing activ-
ity when the oxygen level decreased to 3.0 mg L−1 (40% saturation) in
34.7 g (173 mm) African catfish. In the present study the oxygen level
never reached a critical value of 3 mg L− 1, but decreasing DO levels
where already shown in phase III (Fig. 4) near 7 to 6 mg L−1 in the C.
gariepinus unit. This agrees to findings of Babiker (1979), who described
an increase of the aquatic (branchial) respiratory frequency from 7 mg
L−1 to the highest activity of about 4.0–3.5 mg L−1 oxygen. However,
air breathing in C. gariepinus was described as a facultative mode
(Babiker, 1979; McKenzie et al., 2015), both aquatic respiration and ae-
rial respiration in C. gariepinus seemed to be existent in well oxygenated
water (Moussa, 1957) as C. gariepinus is on the edge of being an obligate
air-breather (Belão et al., 2011). In our study, mainly aquatic respiration
was recorded for C. gariepinus with a regular air breathing activity. How-
ever, caused by reduced water temperatures, the need for oxygen due to
metabolic processes was generally reduced in both fish species. Optimal
metabolic temperatures are in a similar range with 28–32 °C for O.
niloticus (Workagegn, 2012) and 27.5–32.5 °C for C. gariepinus
(Verreth & Den Bieman, 1987; Hogendoorn et al., 1983). Consequently,
the oxygen uptake through the air breathing activity of C. gariepinus is
highly useful for aquaponic cultivation even under saturated oxygen
conditions and supporting system stability.
Oxygen was utilised to identify the maximum fish feed load in the
present aquaponics. Under increasing feed input levels, significant
lower DO was recorded at the beginning of the accumulation phase
(phase III, Table 3, Fig. 4). At this time (days 27–28), the feed input in-
creased from 150 to 200 g d−1. Consequently, the steady phase (maxi-
mum daily feed input) in the present aquaponics was identified at a
feed input of 150 g d− 1 and 10 m2 herbs hydroponics under winter
time conditions. Interestingly, this is in accordance to the successful
use of his UVI system by Rakocy (2012), who estimated the maximum
feed per day and planting area (m²) for gravel and NFT aquaponics in
comparison to a raft aquaponic system. He estimated in his rule of the
thumb a daily feed input of 60–100g in a raft staggered tilapia
aquaponics (steady phase see above) per m2. The gravel and NFT
aquaponics was able to cope with only 25% of that feed input, only
15–25 g m−2 (Rakocy, 2012). This refers to 150–250 g max. feed input
in our aquaponics with 10 m2 gravel bed hydroponics, making parame-
ters of these independent systems comparable. However, the slight
changes of the system design compared with Palm et al. (2014a) already
resulted in a 25% (50 g) less possible feed input (150 vs 200 g) to main-
tain system stability. This is astonishing because the gravel bed has a
large surface area, was well irrigated and increased 20% in size. Addition
of one plant box (120 L water volume) resulted in a total unit water vol-
ume of 3.81 m³ (former 3.75 m³) with a difference of 1.58%. The gravel
substrate volume of 2000 kg per unit used in Palm et al. (2014a) was the
same, but separated into 5 instead of 4 plant boxes in the present study.
 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63 61

This should increase the systems capability to cope with a higher and
not a less nutrient load. However, in the present experiment, we re-
moved the 60 L biofilter. It is obvious that a biofilter inside an
aquaponics system has significant positive influences the general sys-
tem performance even under presence of a gravel bed.
Another important question is the relationship between fish
tank, water volume and optimal hydroponic size. According to
Rakocy (2012), a component ratio of 1 m−3 fish tank allows a volume
of 2 m−3 pea gravel (3 to 6 cm). We used 1.8 m3 fish tank volume and
1.5 m3 gravel. Consequently, our hydroponics could have a 2.4 times larg-
er size to fulfil the recommendation by Rakocy. This might explain that his
feed input range estimated from his rule of thumb of 150–250 g for our
aquaponics was higher (max. 250 g according to Rakocy, 2012) than we
recorded for our unit (150 g) in the present study, suggesting that an in-
crease in gravel bed size × 2.4 would allow only a feed increase of nearly
40% without using a biofilter to provide system stability. Fig. 6. Aquaponic Growth Factor (AGF) as growth differences of aquaponical cultivated
species marked as the deviation from the ideal growth factor (DFIGF) in relation of the
4.4. Biomass development under aquaponic cultivation final biomasses in fish (Factor I = O. niloticus, Factor II = C. gariepinus) to the final
biomass of associated herbs (basil, parsley) in two identical aquaponic units.

Comparison of the two fish species and different plants demonstrat-

ed a disparate productivity under the same system design. Growth of O.
niloticus fingerlings was reduced in comparison to C. gariepinus but
plants performed better yields with O. niloticus (Table 1). Generally,
temperature optima of fish and plants differed under the present exper-
imental conditions. For O. basilicum and P. crispum temperatures were
optimal below 25 °C (Somerville et al., 2014), for O. niloticus and C.
gariepinus optimum temperature was reduced around 17.86% if 28 °C
was used, a level in the range of optimum temperature described by
Workagegn (2012) and Verreth & Den Bieman (1987), respectively. Re-
duced temperature in the present study at 23 °C influenced negatively
the nitrogen excretion rate of fish with generally low growth of plants.
For O. niloticus, linear reduced amounts of ammonia and total waste ni-
trogen was described by McKenzie et al. (1996) between 16, 23 and
33 °C and for C. gariepinus by Buttle et al. (1996) with reduced levels
of total ammonia and total nitrogen at 20 °C and 25 °C (not significant)
and 30 °C with a feed of 40% protein. However, the Aquaponic Growth
Factors (AGF) illustrates the better performance of C. gariepinus (0.41)
compared with O. niloticus (0.29, Fig. 6) using an identical feeding re-
gime. The AGFfish Factor I for fish growth of O. niloticus was calculated
with the final fish biomass of unit I (20,966.7, see Table 1) × final fish
biomass of unit II (29,573.7)− 1 = 0.708–0.71 − DFIGF (= 1) =
|−0.29| with the absolute value of 0.29 for Nile tilapia. The AGFfish Fac-
tor II for C. gariepinus was calculated with final fish biomass of unit II
(29,573.7) × final fish biomass of unit I (20,966.7)−1 = 1.41 − DFIGF
(1) resulted in 0.41 as shown in Fig. 6.
The AGF resulted in better plant performance in combination with O.
niloticus (1.50 in basil, 1.04 in parsley) while plant biomass production
in combination with C. gariepinus was reduced (0.60 in basil, 0.51 in pars-
ley, Fig. 6). For the aquaponic unit I with O. niloticus the AGFplant Factor I
was calculated with the plant biomass of basil (0.30) and parsley (0.55)
divided by plant biomass of unit II with basil (0.12) and parsley (0.27)
with 2.5 for basil subtracted by DFIGF (1) = 1.5 AGFbasil and for parsley;
2.03 (−DFIGF 1) resulted in ~1.04 AGFparsley for O. niloticus (Table 1, Fig.
6). The opposite calculation was done with biomasses for aquaponic
unit II and C. gariepinus. AGFplant Factor II was calculated with the plant
biomass of basil (0.12) and parsley (0.27) divided by plant biomass of
unit I with basil (0.30) and parsley (0.55), resulted in 0.40 for basil
(−DFIGF 1) = |−0.60| AGFbasil and 0.49 (−DFIGF 1) ~ |−0.51| AGFparsley
for unit II with C. gariepinus (Fig. 6). The Aquaponic growth Factor (AGF) is
useful to compare biomass developments in similar experimental de-
signs. A similar result was achieved by Palm et al. (2014b), where C.
gariepinus grew better than O. niloticus and plant yield (lettuce, cucumber,
tomato, basil, Fig. 7) was better in combination with the Nile tilapia but
with opposite fish species stocking in units I & II. As seen in the DFIGF of
fish and plants in the present study (Fig. 6), aquaponic unit II with C.
gariepinus was slightly reduced in nitrogen, resulting in poorer growth
of cultivated herbs. For better plant growth in aquaponic unit I, with O.
niloticus, the retention of nitrate by the plants had to be stronger, as ob-
served by Palm et al. (2014b) with a significant difference of 9.48 mg
L−1 between the two aquaponic units.
We suggest that the observed differences are caused by a different
fish species digestion physiology. Oreochromis niloticus was described
as an opportunistic omnivorous species and has a strong tendency to
herbivory (Lim & Webster, 2006). The African catfish (Clarias gariepinus)
was classified as omnivorous and predacious (Graaf & Janssen, 1996).
Clarias gariepinus is known to be a better feed converter with a more
rapid growth rate than O. niloticus (Haylor, 1989) as found in our
study. The different evolutionary origin of fish has influenced the com-
position of faecal losses in respect to nitrogen and phosphorus as report-
ed by Schneider et al. (2005). Digestibility of nutrient matter was
different in O. niloticus and C. gariepinus. Degani & Revach (1991)
found a better digestibility of protein by tilapia hybrids (O. aureus × O.
niloticus) in contrast to C. gariepinus with poultry meal as the main pro-
tein source. No significant difference in digestibility was observed if
fishmeal was used. Lipid digestion was generally lower in tilapia hybrids
compared to C. gariepinus and carbohydrate utilisation was found to be
more efficient in tilapia. Another study showed a lower protein diges-
tion capability of C. gariepinus if plants (soybean) as sources were used
but it can utilise also diets with a low percentage of animal protein
(Degani et al. 1989). The feed used in this study was designed for omniv-
orous fish like tilapia with different protein fractions of plant, fish and
Fig. 7. Aquaponic Growth Factor (AGF), data adopted from Palm et al. (2014b), as growth
differences of aquaponical cultivated species marked as the deviation from the ideal
growth factor (DFIGF) in relation of the final biomasses in fish (Factor I = C. gariepinus,
Factor II = O. niloticus) to the final biomasses of associated plants (lettuce, cucumber,
tomato, basil) in two identical aquaponic units.
62 U. Knaus, H.W. Palm / Aquaculture 466 (2017) 51–63
animal origin (Aller Aqua 37/10 Float). Protein conversion was better in
C. gariepinus with a higher assimilation of phosphorus and nitrogen and
subsequent lower excretion rate of NH3-N (Table 2) in combination with
reduced faecal losses of nitrogen and phosphorus. The quantitative nitri-
fication process in aquaponic unit II was lower with smaller amounts of
total nitrate in the hydroponic unit and a lower yield of cultured herbs.
Phosphate, as a main nutrient for plant growth, was generally dimin-
ished in contrast to the aquaponic unit with O. niloticus, but could not
had affected plant yield strongly. In contrast to Palm et al. (2014b), phos-
phorus was more than two times higher.
5. Conclusion
The present study demonstrates that depending on the fish species,
the cultivation of herbs in two identical coupled warmwater ebb-and-
flood gravel substrate aquaponic units differed. Plant growth was
under expectation caused by reduced light illumination during winter-
time. O. niloticus had a lower biomass but combined with a better yield
of basil (Ocimum basilicum) and parsley (Petroselinum crispum) com-
pared with C. gariepinus. We suggest the different fish physiology and
a different feed digestibility was responsible for these differences. The
tested aquaponic units performed stable under a maximum feed input
of 150 g day−1, a value corresponding to earlier experiments of gravel
bed aquaponics with similar sized fish tank volumes. Removal of the
biofilter (60 L Biocarrier granules) and the addition of one plant box
(20%) resulted in a reduction of the maximum feed input level of 50 g
per day (25%) compared with an earlier system design. Consequently,
also gravel bed aquaponics require a biofilter if the system is trimmed
for a maximum fish yield. The minimum values of DO were recorded
from the hydroponics, with similar water parameters due to the parallel
hydroponics arrangement. A newly calculated Aquaponic Growth Fac-
tor (AGF) can illustrate differences in fish and plant performances, en-
abling a better understanding of the best possible fish and plant
combinations.
Acknowledgements
We thank the Ministry of Agriculture, Environment and Consumer
Protection of Mecklenburg Western Pomerania for supporting research
in aquaponic fish and plant production. This project was funded
through the pilot project “FishGlassHouse: Innovationsinitiative zur
ressourceneffizienten Nahrungsmittelproduktion in MV” (European
Fisheries Fund-EFF).
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