Agricultural Water Management 71 (2005) 145–166

www.elsevier.com/locate/agwat

Water quality assessment of different reservoir

types in relation to nutrient solution
use in hydroponics
Dietmar Schwarza,*, Rita Groscha, Wolfgang Grossb,1,
Sabine Hoffmann-Hergartenc
a
Institute for Vegetable and Ornamental Crops, Großbeeren/Erfurt e.V.,
Theodor Echtermeyer Weg 1, D-14979 Großbeeren, Germany
b
Institute of Biology, Freie Universität Berlin, Königin Luise Str. 12–16, D-14195 Berlin, Germany
c
Institute for Plant Diseases, Rheinische Friedrich-Wilhelms-Universität Bonn,
Nußallee 9, D-53115 Bonn, Germany
Accepted 14 July 2004

Abstract

Hydroponics requires good quality water. For this purpose, water quality is based on concentra-
tions of specific ions and phytotoxic substances as well as the presence of organisms and substances
that can clog irrigation systems. Here, four irrigation reservoirs, i.e. two rainwater ponds, a peat ditch,
and a natural lake, were analyzed to determine whether or not they conform to water quality
guidelines. Based on our data, the four reservoirs could be divided into two categories in respect to
their water quality. The two rainwater ponds belong to the category characterized by low input of
ionic strength (480 mmol m
1), low concentration of unwanted ions, such as SO42
(63 mmol l
1)
and Zn2+ (3.9 mmol l
1), a moderate bacterial population (lg 4.9 CFU m
1), and moderate algae
density (lg 6.0 cells ml
1). The rainwater ponds were found to contain a good diversity in bacteria (45
species from 25 genera), and a poor diversity of algae (15 species from 4 groups). The other category,
to which the peat ditch and natural lake belong, is characterized by a high ionic strength
(12,200 mmol l
1), high concentrations of alkali ions (Mg2+: 890 mmol l
1; Ca2+: 3.260 mmol l
1;
1; K+: 470 mmol l
1), a moderate bacterial (lg 4.7 CFU ml
1), but low algae density (lg 5.0
cells ml
1). In comparison to the first category, the diversity of the bacteria was poor (seven species

* Corresponding author. Tel.: +49 33701 78206; fax: +49 33701 55391.
E-mail address: schwarz@igzev.de (D. Schwarz).
1
Deceased.

0378-3774/$ – see front matter # 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.agwat.2004.07.005
146 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

from three genera). However, in sharp contrast was the rich algal community detected in the peat
ditch, for which 32 species from six groups were found, whereas in the natural lake, only one group
with seven species was identified. In all reservoirs, species of the genera Paenibacillus and Bacillus
were detected, and small green algae, e.g. Scenedesmus spp., also dominated in each case. Overall,
the bacterial and algal densities showed wide fluctuations between water sources, and neither caused
filter clogging as observed in investigations of others. The quality of the rainwater investigated was
assessed to be well suited for use in hydroponics due to appropriate nutrient concentration (except
Zn2+ in one pond), and lack of potential bacterial and algal development. However, we recommend
water from the natural lake and the peat ditch to be used with care because of the high nutrient
concentration.
# 2004 Elsevier B.V. All rights reserved.

Keywords: Algae; Anions; Bacteria; Cations; Electrical conductivity; Irrigation; Nutrient concentration;
Temperature

1. Introduction

Water quantity and quality are key factors for the cultivation of crops in hydroponic
systems with recirculating nutrient solution. In general, water quality is a complex concept.
However, compared to hydrological definitions, water quality in hydroponics can be
limited to the concentrations of specific ions and phytotoxic substances relevant for plant
nutrition as well as the presence of organisms and/or substances that can clog the irrigation
systems (Tognoni et al., 1998).
Water with low ionic and carbonate concentrations is optimal to avoid salt enrichment
and toxicity in substrates (Schröder and Lieth, 2002). Several publications, in particular for
extension services, provide thresholds or ranges for good, middle, or bad suitability of
water for hydroponic systems expressed in terms of electrical conductivity (EC) or total
salt concentration (Göhler and Drews, 1989; Sonneveld et al., 1991; Ayers and Westcot,
1994; Göhler and Molitor, 2002). The current Dutch norm for water quality for production
in greenhouses is the most precise and defines three classes of water quality based on range
and thresholds for different ions (De Kreij et al., 1999).
Investigations of irrigation water quality have focused mainly on chemical assessments
(Abbas et al., 1993; Avila and Alarcon, 2003; Bini and Bresolin, 1998; Jonnalagadda et al.,
1994; Stolk, 2001), but rarely on pathogenic organisms (Rattink and Van der Sar, 1990;
Berkelmann, 1992). However, water may contain risky microorganisms, such as plant
pathogens (Stanghellini and Rassmussen, 1994; Vanachter et al., 1983; Van Assche and
Vangheel, 1994) or toxic algae (Borowitzka, 1995), for cultivation of many crops. In fungal
species, most of the research performed has been in relation to plant diseases (Hockenhull
and Funck-Jensen, 1983; Rattink and Postma, 1995). The occurrence of other pathogens in
rainwater ponds has also been investigated. Rattink (1991) found Fusarium oxysporum f.
sp. radicis-lycopersici in 78% of rainwater ponds investigated in The Netherlands. Further,
previous studies highlighted some irrigation ponds as a main source of pathogens, such as
Rhizoctonia solani and Pythium spp. Contamination of rainwater ponds occurs by spores of
pathogens from infected air or dust (Gill, 1970; Shokes and McCarter, 1979). Additionally,
algae and cyanobacteria are also able to decrease plant growth due to the release of either
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 147

toxic substances, such as nodularin, goniautoxin, saxitoxin, okadoic acid, or ciguatoxin

(Borowitzka, 1995), or growth regulators, such as abscisic acid or ethylene (Van Staden,
1999). Moreover, microorganisms and algal communities as well as rotifers can lead to
clogging of tubing, filters, and drippers (Juanico et al., 1995). Also, given the right
temperature and nutritional conditions, algae can produce high amounts of mucilage in a
short time, which can result in clogging (Ravina et al., 1997).
On the other hand, certain microorganisms and algae can be beneficial. Some of them
show plant growth promoting effects, like Pseudomonas spp. (Van Peer and Schippers,
1989) and Scenedesmus spp. (Mazur et al., 2001). Others have antagonistic effects against
pathogens, like Bacillus spp. or Pseudomonas spp. (Lemanceau and Alabouvette, 1991;
Grosch et al., 2001) as well as various cyanobacteria (Kulik, 1995; Piccardi et al., 2001).
Additionally, the bacterial population in irrigation water can influence diversity and density
of the microbial population in hydroponics as well as the bacterial population on the root
surface, especially of young plants.
For plant production in hydroponics, growers use water sources with different origins, such
as surface water (lakes, natural and artificial ponds), groundwater (wells), municipal tap water,
rainwater and mixtures of these. Water quality depends mainly on the source that is used. As
rainwater has low ionic strength and usually low microorganism and algal densities, it
conforms to water quality guidelines and is often better than other sources (Sonneveld et al.,
1991). Therefore, a common practice is the collection of rainwater from greenhouse roofs into
ponds. However, as these ponds are fed by atmospheric precipitation, they are vulnerable to
changes in the environment, e.g. eutrophication and acidification. An additional drawback is
that rainwater is not always available for irrigation use due to technical problems in collection
and storage. Therefore, the irrigator must find other water sources, e.g. river or lake; however,
in many cases, such water sources are polluted (e.g. Aboal et al., 1996).
Our objective was to chemically and biologically assess for 1 year two main water
sources used for hydroponic systems, i.e. rainwater and surface water. Therefore, our
investigations focused on characteristics important for hydroponics, i.e. nutrient
concentration, density and diversity of bacterial population, and algal development.

2. Materials and methods

2.1. Description of the water reservoirs

Two artificial rainwater ponds, a peat ditch, and a natural lake were selected for
investigations in 1997 (Table 1). The rainwater ponds (R1: 400 m3, R2: 180 m3) and the
peat ditch (PD) were located at the Institute of Vegetable and Ornamental Crops at
Großbeeren, Germany (lat. 528N). The natural lake (NL) was located at Schwante, about
45 km north of the other sampling sites.

2.2. Sampling and measurement procedure

From 25 February 1997 until 6 January 1998, 1 l water samples were collected every 3
weeks although not always for all locations on each sample date (Table 1). To test for water
148 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Table 1
Characteristics of the four water reservoirs (two rainwater ponds: R1, R2; a peat ditch: PD; a natural lake: NL)
sampled
Characteristic R1 R2 PD NL
Size (m  m  m) 12  25  1.8 8  18  2.4 24  55  2.5 25 ha
Maximum water content (m3) 400 180 2000 500000a
Water influx (m3 year
1) 900 710 661b 125250b
Water efflux (m3 year
1) 557 512 937c 177500c
Greenhouse area (m2) 1500 1000 – –
Sampling dates 9, 12, 18, 21, 9, 12, 18, 21, 24, 12, 18, 21, 21, 24, 30,
(week of the year) 24, 30, 36, 39 27, 30, 36, 39, 24, 30, 36, 39, 36, 42d
42, 46, 49, 54 42, 46, 49, 54
a
Estimated based on a mean depth of 2 m.
b
Based only on precipitation.
c
Based only on evaporation.
d
Ion analyses also at weeks 27, 39, 46.

quality differences inside a single reservoir itself, we took 24 samples from 12 horizontal
and 2 vertical positions of the pond R1 on 25 February. The horizontal sampling points
were distributed net-like, and the vertical points were at the top and 0.2 m above the bottom
of the pond. On the other dates, two samples were taken from each pond, one from the top
and one from the bottom, and one sample was taken from NL. Each sample was separated
into three subsamples for analyses of nutrient concentration (mmol l
1), number of bacteria
(lg CFU ml
1), and determination of the algal community (lg cells ml
1).
Water temperature was measured continuously at two vertical points in the middle of R1
(Pt 100, Messtechnik Geraberg GmbH, Germany) and manually in NL. Total solar
radiation (13G 134, Kriwan Company, Forchtenberg, Germany), humidity (Vaisala HMP
35 A, Helsinki, Finland), and air temperature (DB 2 Kriwan Company, Forchtenberg,
Germany) were monitored outside the reservoir at 2 m height. Precipitation
(501 mm year
1) and evaporation (710 mm year
1) were also recorded, and were
measured hourly and integrated over the day. Water influx and efflux were estimated
corresponding to precipitation and evaporation, respectively. For rainwater ponds, water
collected from the greenhouse roofs was added to the influx and water drawn from the
ponds was added to the efflux.

2.3. Analyses

2.3.1. Chemical analyses

All samples were analyzed for important essential plant nutrients. Total NO3
, NH4+,
PO43
, Mg2+, and SO42
concentrations were measured using spectral photometry (EPOS
analyzer 5060, Eppendorf). Atomic absorption spectral photometry (AAS Vario 6,
Analytik Jena) was used to detect K+, Fe2+, and Zn2+ analysis. Ca2+ was determined by
Titriplex-III solution titration, in the presence of Kali alkaline and Calconcarbon acid. H+
and HCO3
concentrations (mol l
1) were calculated based on pH and concentration of
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 149

Ca2+ and Mg2+ (Manahan, 1979):

½Hþ  ¼ 10
½pH
KðS þ ½Ca2þ þ Mg2þ Þ
½HCO

3¼
½Hþ  þ K
where K = 4.45  10
7, the first ionization constant of H2CO3 and S = 1.028  10
5 M is
the solubility of CO2 in water at 25 8C at 1 atm (air) with a CO2 concentration of 380 ppm.
Electrical conductivity as a measure of total salt concentration (EC, mS m
1, 25 8C) and
pH were measured. The ionic strength (I, mmol l
1) was calculated in the usual manner
(Worch, 1997):
X
I ¼ 0:5 cmi z2i
i

where c is the concentration and z the charge of the ion, i.

2.3.2. Biological analyses

The bacterial population was measured by conventional dilution-plating technique.
Serial dilutions were plated on tryptic soy agar (TSA, VWR International 1.05458) to
quantify the density of the dominant heterotrophic bacterial populations. For Pseudomonas
spp., the selective King’s B medium (KB, VWR International 1.10991) was used. On Petri
dishes, 100 ml of undiluted or diluted rainwater samples was plated in quadruplicate. After
incubation at 25 8C for 48 h, bacterial cells were counted and the decadal logarithm of
colony forming units (lg CFU ml
1) was calculated.
To establish pure cultures for characterization of the individual bacterial strains, one
morphologically representative of each bacterial colony was transferred from the assessed
Petri dishes to a fresh TSA or KB plate. Each strain was identified by analysis of fatty acid
methyl-esters (FAMEs) of total cellular fatty acids in accordance to the method of Sasser
(1990). For fatty acid analysis, bacterial strains were grown on tryptic soy agar (TSBA) for
24 h, and their fatty acids extracted as described by McInroy and Kloepper (1995). TSBA
was prepared in a 2 l flask containing 30 g trypticase soy broth (BBL 11768), 15 g
granulated agar (BBL 11849), and 1 l distilled water. FAMEs were analyzed with a
Hewlett-Packard gas chromatograph model 5890 using a 50 m HP-5 (phenyl silicone)
capillary column (Hewlett-Packard, Newark, DE, USA). Hydrogen was used as the carrier
gas and injections were made in a splitless mode. FAME peaks were analyzed by Microbial
Identification System software (MIS), and bacterial isolates were identified using the MIS
‘Aerobic Library’ (Version 3.7, Newark, DE, USA). Strains with a similarity index below
0.1 were considered as unidentified and those above 0.5 as corresponding well to a
reference strain.
For identification of bacteria strains in the rainwater ponds (R1 and R2), samples were
collected on 25 February (first date) and 2 July. For NL, only on one date (12 June) were
individual bacteria strains determined.
The algal community was assessed by light microscopy, species were enumerated using
a NEUBAUER-counting chamber. Density was assessed in a scale value from 0 to 4, where
0 is 0, 1 is 1–50, 2 is 51–200, 3 is 201–500, and 4 is >500 cells ml
1.
150 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

2.4. Statistics

Results were analyzed by analysis of variance and mean values were compared by
Student’s t-test at P = 0.05. Relationships between anion and cation concentration,
ionic strength, and EC were tested by regression analysis at P = 0.05. Influence of air
temperature, water influx, water efflux, and EC were tested on single nutrient concentration,
EC, pH, bacterial, and algal density by multiple regression analysis at P = 0.05.

3. Results

3.1. Climate conditions

Daily solar radiation fluctuated between 2 MJ m
2 in December and 21 MJ m
2 in

June, and had a mean of 10.6 MJ m
2 (Fig. 1A). The evaporation curve followed the PPFD
curve with a mean daily maximum of 3.8 mm. Evaporation was lowest in December with
0.3 mm. The highest precipitation occurred during the July period at 4.7 mm day
1.
Precipitation was lowest in April and August at 0.5 mm day
1. Daily maximum water
temperature of the ponds was 22.4 8C, 2.6 8C above air temperature (Fig. 1B). While in
January/February, the difference between top and bottom temperature was about 3 8C, and
it was less than 1 8C during the following month until August when differences were no

Fig. 1. Climate conditions at Großbeeren, Germany, during 1997 depicted as mean values between two sampling
dates. (A) Photosynthetic photon flux density (PPFD), evaporation, and precipitation. (B) Air, water temperature,
and humidity.
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 151

Table 2
Chemical composition of water for four water reservoirs (two rainwater ponds: R1, R2; a peat ditch: PD; a natural
lake: NL) sampled
Ion (mmol l
1) Mean Threshold
R1 R2 PD NL
Ca2+ 46.5 34.0 ba 129 300 b 3863 710 a 2656 657 a 2000
Mg2+ 41.8 31.4 c 57.6 29.2 c 1035 87.9 a 740 0b 500
K+ 36.4 56.8 b 33.7 64.1 b 510 98.7 a 429 558 a >b
NH4+ 33.2 35.1 a 6.90 13.7 b 11.4 19.5 b 46.0 50.0 a >
Fe2+ 7.31 19.9 a 1.25 0.76 a 1.12 0.66a 6.86 10.5 a 10
Zn2+ 6.71 3.95 a 1.13 1.07 c 0.31 0.18c 3.52 4.84 b 5
H+c 0.16 0.17 a 0.17 0.22 a 0.02 0.02b 0.02 0.02 b
SO42
52.1 0c 73.9 70.7 c 2954 686 a 1283 684 b 500
PO43
0.94 0.73 b 1.05 0.80 b 0.84 0.5 b 2.98 3.95 a >
NO3
21.4 18.0 a 20.2 18.6 a 9.76 32.3a 11.9 18.6 a >
HCO3
c 61.1 43.6 c 162 351 c 3940 927 a 2767 1056 b 10000
Ionic strength (mmol l
1) 0.347 0.13 c 0.616 1.04 c 16.3 2.8 a 9.27 3.4 b
EC (mS m
1) 3.07 1.38 c 5.51 8.85 c 131 5.83 a 72 18.8 b 50
8dH 0.19 0.13 c 0.42 0.64 c 13.4 2.41 a 9.18 2.24 b
pH 7.48 1.15 a 7.57 1.18 a 7.84 0.36a 7.78 0.44 a
Thresholds of optimal values for good water quality for closed hydroponic systems are given in the last column
(Sonneveld et al., 1991).
a
Different letters depict significant differences among reservoirs at P = 0.05.
b
No thresholds, typically lower than required by plants.
c
Concentrations of H+ and HCO3
calculated.

longer observed. Relative humidity decreased during the summer to the lowest daily
mean of 70%, while it was highest at the end of the year in December with a daily mean of
93%.

3.2. Ion concentration

Based on the results of EC measurements, the water reservoirs can be separated into two
categories. PD and NL exhibited a mean EC of 131 and 72 mS m
1, while R1 and R2 had
much lower EC values of 5.5 and 3.1 mS m
1 (Table 2, Fig. 3A). Similar differences
between the water reservoirs were obtained for the ionic strength. Only in June and at
beginning of July, did EC increase in R1 (in the top region) up to 6.0 mS m
1 and in R2 (in
the bottom region) up to 38 mS m
1, whereas in NL, EC decreased to 42 mS m
1. The
results of the two sampling locations, top and bottom, did not differ significantly. A linear
relationship was confirmed between ionic strength and EC measured with a regression
coefficient of 7.6 (Fig. 2A). The regression coefficient was 10.4 when concentrations of all
ions were added up on a molar basis (data not shown). Anion charge concentration
was larger than cation charge concentration at a ratio of 1.1 (Fig. 2B). Total anion and
cation concentration, EC, and pH correlated with air temperature, water influx, and efflux
(Table 6).
Of all the cations, Ca2+, Mg2+, and K+ displayed the highest concentrations (Fig. 3B).
The concentrations of these ions were significantly higher in PD compared to NL and both
152 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Fig. 2. Relationships between (A) ionic strength (I) and electrical conductivity measured (ECmeas) and (B) anions
and cations (charge concentrations in mmol l
1). Measurements are from water analyses of four water reservoirs
sampled.

rainwater ponds. Maximum values for Ca2+ was 4865 mmol l
1 and for Mg2+
1193 mmol l
1 in PD, and for K+ 1183 mmol l
1 in NL, although the mean value was
also highest in PD. During summer (June) at days 162 and 181, concentrations in the
rainwater ponds reached maximum values, but remained distinctly below concentrations in
the two other reservoirs. For NL and PD, several peaks were observed during the year, but a
significant decrease in concentration of about 50% compared to the mean value was
observed at days 203 and 223 (July, August). The relatively high concentration of Mg2+ and
Ca2+ along with their large contribution to the total ion concentration are responsible for
the high concentration of HCO3
, and thus a pH always above 6 (Fig. 3A). No differences
in pH were found between the ponds. However, the variation in pH was particularly high in
the rainwater ponds (Table 2). Here, a low pH of 6.4 was measured both at the beginning of
the year and on different dates during summer. High pH values (up to 9.5) were detected
only during the summer.
Among the anions, the concentrations of both HCO3
and SO42
were high (Table 2,
Fig. 3). Both showed peaks during the year similar to those described for cations. Most of
the other ions, which are important in plant nutrition as either macronutrients (e.g. NH4+,
NO3
, PO43
) or micronutrients (Fe2+ and Zn2+) were present at concentrations below
50 mmol l
1 (Table 2, Fig. 3). The mean NH4+ concentration in NL was significantly above
mean values in R2 and PD, and showed a peak of 143 mmol l
1 during July. The mean Zn2+
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166
Fig. 3. Time courses of ionic concentration for the four water reservoirs (two rainwater ponds: R1, R2; a peat ditch: PD; a natural lake: NL) sampled. (A) Water electrical
conductivity (EC), pH, and anion concentration (NO3
, PO43
, SO42
) and (B) cation concentration (Ca2+, Mg2+, K+, NH+, Fe2+, Zn2+).

153
154 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Fig. 3. (Continued ).
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 155

concentration in R1 was 6.7 mmol l
1 and this was significantly lower in the other
reservoirs.
K+, Ca2+, and SO42
concentrations correlated with air temperature, water influx, and
efflux (Table 6). Based on simple correlation coefficients, the three factors explained the
variability in ion concentration in equal shares.

3.3. Bacteria

3.3.1. Total bacterial density

The vertical or horizontal sampling position had obviously no effect on bacterial density
except on the first sampling date in February in R1 (data not shown). A mean density of
5.7 lg CFU ml
1 on top and 5.2 lg CFU ml
1 on bottom was estimated on KB and 4.6 and
4.3 lg CFU ml
1 on TSA, respectively. The bacterial density did not differ significantly
between the ponds and varied from 2.7 to 5.6 lg CFU ml
1 on KB and from 1.7 to
4.7 lg CFU ml
1 on TSA (Fig. 4A and B). A high bacterial density above 4 lg CFU ml
1
was measured in both ponds in February, and this decreased by March as assessed on both
media for both ponds. Overall, a significantly higher number of bacteria was counted on the
KB compared to the TSA medium.
The bacterial density measured on both media correlated with air temperature, water
influx, and efflux (Table 6). Based on simple correlation coefficients, influx and efflux
explained the most variability in bacterial density. However, when using the data after the
temperature increase in the middle of March (day 76), temperature exclusively explained
bacterial density (Table 6). A relationship between EC and bacterial density was not found
except with bacterial density on TSA based on results after day 76. Density did not
correlate with single ion concentration, such as NO3
, NH4+, and PO43
, resulting in
R = 0.34 for bacteria on TSA and in R = 0.19 for bacteria on KB.

3.3.2. Identification of bacteria

On the first sampling date in February, 73 bacteria strains (42 from the top and 31 from
the bottom) were identified in R1. From their FAME profiles, these strains were assessed to
comprise 16 genera with 21 species (Table 3). Only nine genera were present both in top

Fig. 4. Time courses of bacterial density for the four water reservoirs (two rainwater ponds: R1, R2; a peat ditch:
PD; a natural lake: NL) sampled. Measurements were carried out on (A) King’s B medium (KB) and (B) tryptic
soy agar (TSA).
156 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Table 3
Identification, isolation frequency (iF), and mean similarity index (S) of bacterial strains in two rainwater ponds
(R1 on two positions, R2 on top) on 25 February 1997 (day 55)
Genera/species R1 top R1 bottom R2 top
iF S iF S iF S
Acinetobacter baumanii 2 0.89 – – – –
A. johnsonii – – 1 0.46 1 0.59
Aeromonas hydrophila – – – – 1 0.61
Agrobacterium radiobacter 1 0.85 – – – –
Arthrobacter atrocyaneus – – 3 0.36 – –
Aureobacterium steroaromaticum 1 0.83 1 0.62 – –
Bacillus cereus 2 0.78 2 0.72 3 0.76
B. megaterium 1 0.72 – – – –
B. pumilus – – – – 1 0.89
B. subtilis – – – – 1 0.66
B. thuringiensis – – 1 0.67 1 0.79
Brevundimonas vescularis 1 0.37 – – – –
Cellumonas flavigena 1 0.70 1 0.71 – –
Chryseobacterium indologeneses 6 0.90 4 0.92 – –
Cytophaga johnsonae – – – – 1 0.40
Hydrogenophaga pseudoflava – – 1 0.27 – –
Janthinobacterium lividium 7 0.77 1 0.39 1 0.82
Micrococcus kristinae – – – – 1 0.69
M. varians 2 0.68 3 0.70 1 0.53
Paenibacillus macerans 4 0.41 10 0.51 2 0.60
Pseudomonas chlororaphis 1 0.83 1 0.88 5 0.80
P. fluorescens 1 0.78 – – – –
P. marginalis – – – – 1 0.87
P. putida – – – – 4 0.72
Rhodococcus globerulus – – 1 0.73 – –
Sphingobacterium heparinum 5 0.37 – – – –
Sp. spiritivorum 4 0.39 – – – –
Stenotrophomonas maltophilia 3 0.68 1 0.67 – –

and bottom samples. The species Janthinobacterium lividium, Sphingobacterium spir-

itivorum, and Sp. spiritivorum were isolated more frequently in top samples, whereas
Paenibacillus macerans occurred more frequently in bottom samples. No clear differences in
frequency of the genera Bacillus, Chryseobacterium, and Micrococcus were observed
between top and bottom samples. In R2, 24 strains were identified in total, which comprised 8
genera with 14 species. Here, Bacillus and Pseudomonas were the most frequent genera, and
four and three species of these genera were isolated, respectively. The genera
Janthinobacterium, Paenibacillus, Bacillus, Micrococcus, Pseudomonas, and Acinetobacter
were identified both in R1 and R2. Frequently isolated genera, such as Janthinobacterium,
Chryseobacterium, Bacillus, Micrococcus, and Pseudomonas, were identified with a
similarity index above 0.5, whereas the coefficients of the genera Sphingobacterium and
Paenibacillus were below 0.5. A similarity index below 0.5 does not allow a clear
identification of these isolates following the definition of the MIS database used.
In the samples collected in summer, a total of 57 bacteria strains were characterized with
FAMEs from R1, R2, and NL corresponding to 34 strains from 18 genera (Table 4). In each
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 157

Table 4
Identification, isolation frequency (iF), and mean similarity index (S) of bacteria strains in two rainwater ponds
(R1, R2) on 12 August 1997 (day 223) and in the natural lake (NL) on 1 July 1997 (day 181)
Genera/species R1 R2 NL
iF S iF S iF S
Acinetobacter radioresistens 1 0.88 – – – –
Aeromonas hydrophila 5 0.85 1 0.86 – –
A. salmonica 2 0.19 – – – –
A. schuberti – – – – 1 0.63
A. sobria – – – – 2 0.62
A. trota 1 0.91 – – – –
A. veronii – – – – 1 0.82
Alcaligenes xylosoxydans 1 0.69 – – –
Arthrobacter ilicis 1 0.10 – – – –
Bacillus cereus 2 0.66 1 0.79 – –
B. coagulans – – 1 0.54 – –
B. licheniformis – – 1 0.95 2 0.78
B. megaterium 1 0.95 – – – –
B. pumilus – – 1 0.94 2 0.80
B. subtilis – – 1 0.82 1 0.83
B. thuringiensis 1 0.70 – – – –
Burkholderia pickettii – – 1 0.74
Cellulomonas flavigena – – 1 0.59
Clavibacter michiganense – – 2 0.86
Comamonas acidovorans 1 0.95 – – – –
Enterobacter taylorae 1 0.92 – – – –
Erwinia carotovora 1 0.67 – – – –
E. chrysanthemi 1 0.87 – – – –
Kluyvera ascorbata – – 1 0.85 – –
K. cryocrescens 1 0.75 1 0.44 – –
Micrococcus halobius – – 1 0.61 – –
M. varians – – 1 0.60 – –
Paenibacillus macerans 2 0.54 1 0.66 2 0.50
Pantoea agglomerans – – 3 0.68
Rhodococcus fascians 1 0.85 – – – –
Staphylococcus epidermidis – – 1 0.74
St. Hominis – – 1 0.46 – –
St. Warneri – – 1 0.78
Stenotrophomonas maltophilia – – 2 0.72

rainwater pond, the bacteria comprised 11 genera. Seven different bacterial strains were
identified from NL, which comprised 3 genera. Species of the genera Aeromonas, Bacillus,
Kluyvera, and the species Paenibacillus macerans occurred in all three reservoirs. Nearly
all genera identified were isolated in low frequencies, except for Aeromonas and Bacillus.

3.4. Algae

The total algal density differed significantly between the reservoirs, and was highest in
R1 followed by R2, PD, and NL (Fig. 5A). The density curve resembles a sinus function
during the 1-year time course with two peaks, one at the beginning of May and the other at
158 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Fig. 5. Time courses of algae occurrence for the four water reservoirs (two rainwater ponds: R1, R2; a peat ditch:
PD; a natural lake: NL) sampled. (A) Total algal density. Dominant species (>100 cells ml
1); (B) Chlorhormi-
dium pseudistichococcus; (C) Haematococcus pluvialis; and (D) Scenedesmus longispina and S. quadricauda.

end of August. In R1, the maximum density was reached at 6.8, in R2 at 6.6, and in PD at
5.5 lg cells ml
1. In NL, the density was measured only in June when it amounted to
2.4 lg cells ml
1. Samples collected near the surface and the bottom of the ponds did not
differ significantly. Algal density correlated with temperature, water influx to the
reservoirs, and EC (Table 6). Based on simple correlation coefficients, EC (
0.65)
explained most of the variability in algal density followed by influx (
0.33) and then
temperature (0.27). However, algal density did not correlate significantly with NO3
and
PO43
, but was negatively correlated with NH4+. The coefficient of determination was low
(R = 0.42) and the simple correlation coefficient for NH4+ was 0.27.
Large differences were found in the algal community among the four reservoirs
investigated. The diversity of the species was rather poor in the rainwater ponds: 10 in R1,
13 in R2, and 7 in NL (Table 5). In PD, however, the algal community was rich with 32
species found. Unicellular green algae dominated throughout the year, mainly consisting of
Chlorhormidium pseudostichococcus (Fig. 5B), Haematococcus pluvialis (early spring,
Fig. 5C), and Scenedesmus spp. (Fig. 5D). The density of these three species correlated
with air temperature, water influx, and efflux (Table 6). Based on simple correlation
coefficients, influx explained the most variability in the density of Chlorhormidium and
Haematococcus, whereas efflux explained the most variability in the density of
Scenedesmus. The time course of the single species was similar to the time course of
the total density described above. Highest cell density of Ch. pseudostichococcus (6.48 lg
cells ml
1) was observed in August, and of H. pluvialis (R1 5.4 lg cells ml
1) and S.
longispina (6.4 lg cells ml
1) in May. The density of H. pluvialis and S. longispina differed
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 159

Table 5
Algae species present in the four water reservoirs (two rainwater ponds: R1, R2; a peat ditch: PD; a natural lake:
NL) sampled
Group Species R1 R2 PD NL
Bacillariophyta Amphipleura pellicula 0.33
Diatoma elongatum 0.17
Navicula spec. 0.04 0.08
Nitzschia acicularis 1.83
Nitzschia sigmoidea 0.25
Rhizosolenia longiseta 0.04
Synedra acus 0.08
Chlorophyta Acanthosphaera zachariasii 2.25
Ankistrodesmus angustus 0.23 0.21
Ankistrodesmus spirilliformis 0.25
Carteria multifilis 0.19
Chlamydomonas reinhardtii 0.60 0.38
Chlorhormidium pseudistichococcus 3.60 0.88
Choricystis minor 1.13 0.69
Crucigenia quadrata 0.25
Dictyoshaeridium spp. 0.17
Haematococcus pluvialis 3.00 1.92
Kirchneriella contorta 0.08
Micractinium pusillum 0.04
Pediastrum integrum 0.17
Pediastrum simplex 0.04 1.00
Raphidium spp. 0.08
Scenedesmus longispina 3.73 3.38 1.50 2.00
Scenedesmus obliquus 1.00
Scenedesmus quadricauda 0.33 0.50 1.29
Selenastrum bibraianum 0.08 0.50
Stichococcus bacillaris 0.46
Tetraedron minimum 0.42 0.50
Tetraedron trigonum 0.08
Tetrastrum glabrum 0.50
Treubaria triappendiculata 1.00
Chrysophyta Dinobryon sociale 0.17
Cryptophyta Chroomonas nordstedtii 0.35
Cryptomonas ovata 0.04
Dinophyta Peridinium tabulatum 0.58
Euglenophyta Euglena acus 0.58
Euglena pisciformis 0.20 0.08
Euglena sanguinea 0.12 0.17
Euglena viridis 0.12
Euglena spp. 0.20 0.27
Phacus occilans 0.07
Phacus triquester 0.04
Trachelomonas euchlora 0.38
Trachelomonas hispida 0.04
Trachelomonas volvocina 0.04
Mean values from all sampling dates (0 is 0, 1 is 1–50, 2 is 51–200, 3 is 201–500, and 4 is >500 cells ml
1).
160 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Table 6
Coefficients of determination (R) and regression coefficients for the influence of mean air temperature (T), water
influx and water efflux, and electrical conductivity measured (EC) for the four reservoirs throughout a year on
characteristics of chemical composition, bacteria measured on tryptic soy agar (TSA, lg CFU ml
1) or King’s B
medium (KB, lg CFU ml
1), and algal community (lg cells ml
1)
Characteristic n R T (8C) Influx (m3) Efflux (m3) EC (mS m
1)
+
1 * *
K (mmol l ) 85 0.612 20.0
0.346
0.589
Ca2+ (mmol l
1) 85 0.654 87.9*
2.79*
3.80*
SO42
(mmol l
1) 85 0.636 71.1*
1.95*
2.86*
Anion (mmol l
1) 69 0.636 234*
7.63*
7.84*
Cation (mmol l
1) 69 0.639 224*
7.12*
7.10*
EC (mS m
1) 69 0.697 3.46*
0.106*
0.117*
pH 69 0.569 0.067*
0.0014* 0.00084
Bacteria TSA 42 0.419 0.034* 0.0030*
0.0014* 0.0043
After day 76 34 0.681 0.059* 0.00011 0.00054 0.006*
Bacteria KB 42 0.555 0.045* 0.0038*
0.0016
0.0016
After day 76 34 0.675 0.69* 0.0019 0.00034 0.0030
Algae 44 0.723 0.06*
0.0019* 0.0013
0.015*
Chlorhormidium spp. 19 0.660 0.176*
0.00076
0.0033*
0.125
Haematococcus spp. 16 0.841 0.114*
0.00049
0.0028*
0.092
Scenedesmus spp. 27 0.862 0.0356
0.0022* 0.0021*
0.016*
Asterisk marks significant differences at P < 0.05.

significantly between the reservoirs and showed the highest values in R1 over most of the
year. S. longispina was the only species found in all four reservoirs, although in PD and NL
only in moderate densities (Table 5). Other species observed at higher densities were
Chlamydomonas reinhardtii (R1), Nitzschia acicularis (PD), and Scenedesmus quad-
ricauda (PD) during April to May as well as Choricystis minor (R1, R2), Stichococcus
bacillaris (PD), and Acanthosphaera zachariasii (PD) during August (Table 5). Overall,
the cell density showed wide fluctuations, especially in summer. Species observed
occasionally and usually with low cell densities are also listed in Table 5. Most of the
species belonged to either Chlorophyta, Euglenophyta, or Bacillariophyta (diatoms).

4. Discussion

4.1. Ions

The most important characteristics of chemical quality of irrigation water are total
soluble salt concentration, concentration of individual ions, the concentration of substances
that may be toxic (e.g. zinc), and the ratio of bicarbonate to calcium and magnesium
(Hanan, 1998).
All results on EC as measured for total soluble salt concentration from the rainwater
ponds (Fig. 3) are below the threshold of 50 mS m
1 reported for best irrigation water used
for hydroponics (De Kreij et al., 1999), or waters regarded as generally safe by the US
Salinity Laboratory (25 mS m
1; Richards, 1954). These values are higher compared to
data from clean areas, such as Zimbabwe, Africa where 0.9–1.4 mS m
1 was measured
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 161

(Jonnalagadda et al., 1994); however, they are within those from coastal areas, such as
Hong Kong, which give values of 1.5–10 mS m
1 (Sequeira and Lai, 1998), or Southern
Brazil with 3–4 mS m
1 (Abbas et al., 1993). Our values are comparable with results from
areas where rainwater is collected for use in horticulture, such as Spain with 1.0–
3.9 mS m
1 (Avila and Alarcon, 2003), Italy with 2.4 mS m
1 (Bini and Bresolin, 1998),
and The Netherlands with 2.5–4.8 mS m
1 (Stolk, 2001, pp. 23–24). Larger inputs come
either from environmental pollution (Hong Kong) or from the sea via NaCl (Southern
Brazil). Our results also correspond to EC of <20 mS m
1 found in reservoirs in Middle
Europe that have low levels of electrolytes (Kalbe, 1997). On the other hand, mean EC of
PD and NL are above the thresholds for best irrigation water, while the EC of NL agrees
with normal EC in surface reservoirs in Middle Europe between 30 and 90 mS m
1, but the
EC of PD is higher (Kalbe, 1997). The mean value is even higher compared to ECs of water
from polluted sources, such as wells (Hanan, 1998) and rivers, e.g. the river Rhine
(100 mS m
1) or the Vaal River in South Africa (76 mS m
1; Roos and Pieterse, 1995).
Seasonal variations in EC and ionic strength, namely the two peaks in June and August/
September, were possibly caused by natural and anthropogenic sources as shown in several
short-term investigations (Abbas et al., 1993; Jordan et al., 1995; Roos and Pieterse, 1995;
Stolk, 2001). Heinen (1997) defines an approximate calculation of EC (dS m
1) to be the
sum of the concentrations of all ions (mmol l
1) times 10. Our data are in good agreement
with this estimation. Additionally, EC/ionic strength ratios (Fig. 2A) also agree with data
from the literature, i.e. McNeal et al. (1970) calculated an insignificantly higher
regression’s coefficient of 7.8 compared to our calculation of 7.6 (Fig. 2A). One reason for
our lower coefficient might be the influence of a different ion composition (McNeal et al.,
1970). Moreover, the amount of total ions present could be underestimated. For example,
many authors have reported NaCl or NaSO4 to be found in surface and rainwater (Avila and
Alarcon, 2003; Roos and Pieterse, 1995; Sequeira and Lai, 1998; Stolk, 2001); however, our
ion measurements did not include Na+ and Cl
. Furthermore, the concentration of HCO3
and
H+ was based on calculations. Thus, particularly additional amounts of Na+ are likely and
those of Cl
and HCO3
are possible. Usually, Na+ concentrations in rainwater are given at
12 mmol l
1 (Jonnalagadda et al., 1994). Rainwater collected near the sea shows higher
concentrations of up to 200 mmol l
1 (Abbas et al., 1993; Stolk, 2001), and in polluted surface
water, it can reach 6000 mmol l
1 (Roos and Pieterse, 1995). The lower cation concentration
compared to anion concentration supports the notion that Na+ was underestimated (Fig. 2B).
Of the calculated anion/cation ratio only that for rainwater ponds (0.84) showed an anion
deficit. For rainwater especially, an anion deficit has often been reported, e.g. by Edmonds
et al. (1995) with 0.9, caused by the portion of organic anions. Anion/cation ratios are
described to vary between 0.2 and 2 depending on climate and influence of pollution or sea
(Avila and Alarcon, 2003; Edmonds et al., 1995; Jonnalagadda et al., 1994).
Normally, only very low concentrations of PO43
, NO3
, and K+ are found in deep wells or
in uncontaminated irrigation water (Hanan, 1998). Except for K+ and PO43
in NL, similar
results were obtained for all four reservoirs. K+, Ca2+, and Mg2+ as well as SO42
and HCO3

were available in high amounts in PD and NL compared to concentrations in rainwater ponds,
while all other ions were within a normal range (Abbas et al., 1993; Avila and Alarcon, 2003;
Stolk, 2001). These high concentrations of SO42
in PD and of NH4+ and PO43
in NL
indicate a trend to eutrophication. Concentrations >3000 mmol l
1 signal pollution by
162 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

SO42
, >1.5 mmol l
1 by PO43
, and >100 mmol l
1 by NH4+ (Hanan, 1998). PD and NL
are surrounded by farmlands, therefore the peaks in NH4+ observed (Fig. 3) may have been
caused by application of fertilizer or manure. However, the high concentrations of NH4+ and
PO43
do not represent a problem in irrigation water although these values are characteristic
for polytrophication (Kalbe, 1997). On the other hand, the sulfate concentration is higher than
the threshold given by Sonneveld et al. (1991) of 500 mmol l
1 for good quality of irrigation
water (Table 2). Additionally, the Ca2+ concentration in PD and NL also exceeds the threshold,
and here it was estimated to be 2000 mmol l
1. The concentrations are two to four times higher
than the average concentration in European surface waters (1000 mmol l
1, Wetzel, 2001).
Ca2+ concentrations varied between 1700 and 4700 mmol l
1, but the range for the Ca/Mg
ratio is within 4–5, which is normal for surface water (Worch, 1997). In the two rainwater
ponds, the ratio was 1.1 (R1) and 2.2 (R2). Jonnalagadda et al. (1994) and Edmonds et al.
(1995) also found ratios below 2 for rainwater at different locations. Dominant ions in our
rainwater were Ca2+ and SO42
, which could account for most of the variation in alkalinity as
has been reported in many other regions in Europe (Avila and Alarcon, 2003; Bini and
Bresolin, 1998; Stolk, 2001).
Among the ions that may be toxic, Zn2+ was found to be above a concentration of
5 mmol l
1, and therefore should be taken into account for the use of irrigation water (De
Kreij et al., 1999). When water is collected from greenhouses made from galvanized
metals, the Zn2+ concentration can be problematic particularly in closed hydroponic
system with recirculation of nutrient solution as indicated by high Zn2+ concentrations in
R1 (Fig. 3B). However, this is well known and taken into consideration by the growers.
Excluding this exception, it can be concluded that rainwater ponds contain water well
suitable for use in hydroponics and for irrigation of plants. Water from PD and NL is only
for limited use due to high SO42
and Ca2+ concentrations.

4.2. Bacteria

In general, counted numbers of bacteria were higher compared to results reported by

others (Berkelmann, 1992; Schwarz et al., 1999). This might be due to the use of different
media and different climate conditions during the time of sampling. Previous investigations
by Berkelmann (1992) were carried out with irrigation water prepared by mixing rainwater
and tap water, where a low bacterial density in the tap water probably diluted the density of
bacteria in the rainwater. Schwarz et al. (1999) observed a similar increase of bacterial
density (Fig. 4B) throughout the year, although on a lower level, which indicates an
influence of radiation or temperature. Our results (Table 6) confirmed temperature to be an
influencing factor. After the second sampling date (day 76), the bacterial density increased
with higher temperature. Therefore, a relationship between replication rate as well as
microbial activity and temperature can be assumed. In the rainwater, bacteria might have
been preserved because of low temperature (<0 8C, Fig. 1). For bacteria, such as
Pseudomonas spp., Bacillus spp., or Staphylococcus spp., the optimal growth temperature
is >15 8C (Palleroni and Doudoroff, 1972; Berkeley et al., 1984). However, the microbial
activity in the water reservoirs was not monitored. Bacterial densities above lg
4.5 CFU ml
1 as monitored on KB media for most of the sampling dates (except in PD)
could possible lead to filter clogging (Ayers and Westcot, 1994).
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 163

The low correlation coefficients calculated for the influence of different factors
on bacterial density (Table 6) indicate that there are other influencing factors which
remain to be identified. One of these factors might be the relation between organic nitrogen
and organic carbon in the water; however, here, the contents were not determined.
Additionally, most of the characterized bacteria need aerobic conditions for optimal
growth. Therefore, the oxygen concentration, affected by temperature, could be another
factor.
Generally, the number of bacteria on TSA was lower compared to the number on KB
although a higher number of heterotrophic bacteria was expected. KB was used especially
for the selection of Pseudomonas spp., which were frequently observed in hydroponics on
different greenhouse crops (Berkelmann, 1992; Waechter-Kristensen et al., 1994).
Pseudomonas spp. were detected on the first sampling date in February both in R1 and R2
(Table 3), but not on the second sampling date in July. How bacteria from rainwater affect
the development of bacterial populations on plants and in nutrient solution in hydroponics
is still not known and remains to be investigated.
The 27 different genera isolated demonstrate that rainwater ponds serve as microbial
habitats for a diverse bacterial microflora. These results provide only an indication of the
diversity, because a high percentage of bacteria do not grow on artificial media in Petri
dishes. Additionally, we can also assume that more genera could be identified when using
different media. Only the genera Paenibacillus and Bacillus were found in ponds on both
dates. Isolates of these genera produce spores and might be more resistant against changes
in environmental conditions. The occurrence of other genera detected seems to be more
accidental and varied from pond to pond as well as over the time. No causal relationship
could be observed.
The results in Table 6 indicate that irrigation water reservoirs are both a source of plant
pathogens, such as Clavibacter michiganense, Erwinia carotovora, and E. chrysanthemi,
and a habitat of plant growth promoting rhizobacteria, such as Pseudomonas fluorescens, P.
putida, Paenibacillus macerans, or Bacillus subtilis. Other bacterial taxa characterized
here, such as Aeromonas spp., Acinetobacter spp., and Stenotrophomonas maltophilia, are
widely distributed in nature (Hazen et al., 1978; Denton and Kerr, 1998).

4.3. Algae

In our studies, planktonic species were almost exclusively monitored due to our
sampling method. Benthic algae or species from the neuston were observed only
occasionally in our samples. In general, the density and diversity of the algal community
was typical for oligo- to slightly eutrophic ponds. The absence of cyanobacteria blooms
was also indicative for moderate eutrophic conditions. Although the nutrient concentration
was lower in the rainwater ponds compared to the other reservoirs, the algal density was
higher (Fig. 5). This is most likely due to the low abundance of predators.
Only one species was found in all four reservoirs, namely S. longispina. This species is
ubiquitous in fresh water. Interestingly, the composition of the species determined in the
rainwater ponds differed from that found in natural lakes under similar climate conditions
(Krienitz et al., 1996). Eight species occurred in both rainwater ponds (Table 5) all of which
are commonly found in this type of environment. In all cases, the development of these
164 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

species differed significantly between both ponds, except for Chlorhormidium

pseudostichococcus and Euglena spp. Most likely this was influenced by specific pond
characteristics. The reservoir with the highest total ion concentration, PD, was also richest
in diversity of algae species. Lewin and Guillard (1963 in Roos and Pieterse, 1995) found a
stimulatory effect on algae growth caused by high Ca concentrations. In PD, the Ca
concentrations were highest (3870 mmol l
1) compared with the other reservoirs.
Chlamydomonas spp., Haematococcus pluvialis, and Scenedesmus spp. are known to
cause growth promoting effects on plants (Ördög, 1999), and thus are beneficial in
hydroponics. On the other hand, Chlamydomonas spp. become dominant in closed
hydroponic systems, and thereby limit growth of other species (Nonomura et al., 2001).
However, we did not observe this in our samples. None of the species identified here are
known to have potential toxic effects on other organisms or with potential for antagonistic
effects on plant pathogens.
In summary, algal species known to produce significant amounts of mucilage were not
observed. Ravina et al. (1997) found mucous organisms (colonial protozoa and to a lesser
extend, also bryozoa) responsible for clogging. The absence of filamentous species as well
as mucilage-producing algae and the relatively low cell densities connected with low
temperature conditions suggest that algae are usually not the main cause of filter or emitter
clogging (Aboal et al., 1996; Juanico et al., 1995; Ravina et al., 1997). Both the density of
bacteria and algae are negatively correlated to some extent to the total nutrient
concentration in the reservoirs (Table 6), but not to any single ion concentration. However,
a higher nutrient concentration seems to enhance the diversity of algae as observed in PD
(Table 5). The tendency to eutrophication and polytrophication in NL reduced the diversity
of both bacteria and algae.

Acknowledgement

The Ministries of Consumer Protection, Food, and Agriculture of the Federal Republic
of Germany, Brandenburg, and Thüringen supported this study.

References

Abbas, M.Z.M., Bruns, R.E., Scarminio, I.S., Ferreira, J.R., 1993. A multivariate statistical analysis of the
composition of rainwater near Cubato, Sp., Brazil. Environ. Pollut. 79, 225–233.
Aboal, M., Prefasi, M., Ascenio, A.D., 1996. The aquatic microphytes and macrophytes of the Transvase Tajo–
Segura irrigation system, southeastern Spain. Hydrobiologia 340, 101–107.
Avila, A., Alarcon, M., 2003. Precipitation chemistry at a rural Mediterranean site: between anthropogenic
pollution and natural emissions. J. Geophys. Res. 108, 4278.
Ayers, R.S., Westcot, D.W., 1994. Water quality for agriculture, third ed. FAO Irrigation and Drainage Paper. Food
and Agriculture Organization of the United Nations Rome.
Berkeley, R.C.W., Logan, N.A., Shute, L.A., Capey, A.G., 1984. Identification of Bacillus species. In: Bergan
(Ed.), Methods in Microbiology, vol. 16. Academic Press, London, pp. 291–328.
Berkelmann, B., 1992. Charakterisierung der Bakterienflora und des antagonistischen Potentials in der zirkulier-
enden Nährlösung einer Tomatenkultur (Lycopersicon esculentum MILL.) in Steinwolle. Ph.D. thesis,
University Göttingen.
 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166 165

Bini, C., Bresolin, F., 1998. Soil acidification by acid rain in forest ecosystems: a case study in northern Italy. Sci.
Total Environ. 222, 1–15.
Borowitzka, M.A., 1995. Micro-algae as sources of pharmaceutical and other biologically active compounds. J.
Appl. Phycol. 7, 3–15.
De Kreij, C., Voogt, W., van den Bos, A.L., Baas, R., 1999. Bemestings adviesbasis substraten. Proefstation voor
Bloemisterij en Glasgroenten. Vestiging Naaldwijk, p. 145.
Denton, M., Kerr, K.G., 1998. Microbiological and clinical aspects of infections associated with Stenotropho-
monas maltophilia. Clin. Microbiol. Rev. 11, 7–80.
Edmonds, R.L., Thomas, T.B., Blew, R.D., 1995. Biogeochemistry of an old-growth forested watershed, Olympic
National Park, Washington. Water Resources Bull. 31 (3), 409–419.
Gill, D.I., 1970. Pathogenic Pythium from irrigation ponds. Plant Dis. Rep. 54, 1077–1079.
Göhler, F., Drews, M., 1989. Hydroponische Verfahren bei der Gemüseproduktion in Gewächshäusern. Land-
wirtschaftsausstellung der DDR, agrabuch, p. 108.
Göhler, F., Molitor, H.-D., 2002. Erdelose Kulturverfahren im Gartenbau. Verlag Eugen Ulmer, p. 267.
Grosch, R., Junge, H., Kofoet, A., 2001. Prüfung von Isolaten der Bacillus—Gruppe gegen Fusarium
oxysporum f. sp. radicis-lycopersici an Tomatenpflanzen in erdeloser Kultur. Gartenbauwissenschaften 66,
262–269.
Hanan, J.J., 1998. Greenhouses. Advanced Technology for Protected Horticulture. CRC Press, Boca Raton, FL.
Hazen, T.C., Fliersmann, C.B., Hirsch, R.P., Esch, G.W., 1978. Prevalence and distribution of Aeromonas
hydrophila in the United States. Appl. Environ. Microbiol. 36, 731–738.
Heinen, M., 1997. Dynamics of water and nutrients in closed, recirculating cropping systems in glasshouse
horticulture. Thesis, Agricultural University Wageningen, The Netherlands, p. 270.
Hockenhull, J., Funck-Jensen, D., 1983. Is damping less of a problem in hydroponics than in traditional growing
systems? Acta Hort. 133, 137–145.
Jonnalagadda, S.B., Makadho, J., Matinde, N., Karimanzirza, R.P., Makarau, A., 1994. Chemical composition of
rainwater and air quality in Zimbabwe, Africa. Sci. Total Environ. 144, 261–271.
Jordan, T.E., Correll, D.L., Weller, D.E., Goff, N.M., 1995. Temporal variation in precipitation chemistry on the
shore of the Chesapeake Bay. Water Air Soil Pollut. 83, 263–284.
Juanico, M., Azov, Y., Teltsch, B., Shelef, G., 1995. Effect of effluent addition to a freshwater reservoir on the filter
clogging capacity of irrigation water. Water Res. 29 (7), 1605–1702.
Kalbe, L., 1997. Limnische Ökologie. B.G. Teubner Verlagsanstalt Stuttgart, Leipzig, 296 pp.
Krienitz, L., Pasparzak, P., Koschel, R., 1996. Long term study on the influence of eutrophication, restoration and
biomanipulation on the structure and development of phytoplankton communities in Feldberger Haussee,
Germany. Hydrobiologia 330, 89–110.
Kulik, M.M., 1995. The potential for using cyanobacteria (blue–green algae) and algae in the biological control of
plant pathogenic bacteria and fungi. Eur. J. Plant Pathol. 101, 585–599.
Lemanceau, P., Alabouvette, C., 1991. Biological control of fusarium diseases by fluorescent Pseudomonas and
non-pathogenic Fusarium. Crop Protection 10, 279–286.
Manahan, S.E., 1979. Environmental Chemistry, third ed. Willard Grant Press, Boston.
Mazur, H., Konop, A., Synak, R., 2001. Indole-3-acetic acid in the culture medium of two axenic green
microalgae. J. Appl. Phycol. 13, 35–42.
McInroy, J.A., Kloepper, W., 1995. Survey of indigenous bacterial endophytes from cotton and sweet corn. Plant
Soil 173, 337–342.
McNeal, B.L., Oster, J.D., Hatcher, J.T., 1970. Calculation of electrical conductivity from solution composition
data as an aid in situ estimation of soil salinity. Soil Sci. 110, 405–414.
Nonomura, T., Matsuda, Y., Bingo, M., Onishi, M., Matsuda, K., Harada, S., Toyoda, H., 2001. Algicidal effect of
3-(3-indolyl)butanoic acid, a control agent of the bacterial wilt pathogen, Ralstonia solanacearum. Crop
Protection 20, 935–939.
Ördög, V., 1999. Beneficial effects of microalgae and cyanobacteria in plant/soil-systems, with special regard to
their auxin- and cytokinin-like activity. In: Proceedings of the International Workshop and Training Course on
Microalgal Biology and Biotechnology. Mosonmagyaròvàr, Hungary, p. 43.
Palleroni, N.J., Doudoroff, M., 1972. Some properties and taxonomic subdivisions of the genus Pseudomonas.
Annu. Rev. Phytopathol. 10, 73–100.
166 D. Schwarz et al. / Agricultural Water Management 71 (2005) 145–166

Piccardi, R., Biondi, N., Frosini, A., Rodolfi, L., Margheri, M.C., Tredici, M.R., 2001. Cyanobacteria of the genus
Nostoc as a source of novel agroactive compounds. In: Proceedings of the International Symposium on
Microalgae and Seaweed Products in Plant/Soil-Systems. Mosonmagyarovar, Hungary, p. 18.
Rattink, H., Postma, J., 1995. Biological control of soil-borne fungi in glasshouse crops on closed systems. In:
Manka (Ed.), Environmental Biotic Factors in Integrated Plant Disease Control. The Polish Phytopathological
Society, Poznan, pp. 455–459.
Rattink, H., Van der Sar, M., 1990. Waterbassins besmet met Fusarium voetziekte. Groenten en Fruit 46, 37.
Rattink, H., 1991. Epidemiology of Fusarium crown and root rot in artificial substrate sytems. Medelingen
Faculteit Landbouwwetenschappen Rijksuniversiteit Gent 56 (2b), 423–430.
Ravina, I., Paz, E., Sofer, Z., Marcu, A., Schischa, A., Sagi, G., Yechialy, Z., Lev, Y., 1997. Control of clogging in
drip irrigation with stored treated municipal sewage effluent.. Agric. Water Manage. 33 (2–3), 127–137.
Richards, L.E., 1954. Diagnosis and improvement of saline and alkali soils. Agric. Handbook No. 60, USDA,
Washington, DC, p. 160.
Roos, J.C., Pieterse, A.J.H., 1995. Salinity and dissolved substances in the Vaal River at Balkfontein, South Africa.
Hydrobiologia 306, 41–51.
Sasser, M., 1990. Identification of bacteria through fatty acid analysis. In: Klement, Z., Rudolph, K., Sands, D.C.
(Eds.), Methods in Phytobacteriology. Akademiai Kiado, Budapest, pp. 199–204.
Schröder, F.-G., Lieth, J.H., 2002. Irrigation control in hydroponics. In: Savvas, D., Passam, H. (Eds.), Hy-
droponic Production of Vegetables and Ornamentals. Embryo Publications, Athens, Greece, pp. 263–298.
Schwarz, D., Ruppel, S., Kuchenbuch, R., 1999. Nitrogen cycle and micro-organisms in a hydroponic system as
influenced by the amount of nitrogen applied. Acta Hort. 481, 371–378.
Sequeira, R., Lai, C.C., 1998. Small-scale spatial variability in the representative ionic composition of rainwater
within urban Hong Kong. Atmos. Environ. 32 (2), 133–144.
Shokes, F.M., McCarter, S.M., 1979. Occurrence, dissemination and survival of plant pathogens in surface
irrigation ponds in southern Georgia. Phytopathology 69, 510–516.
Sonneveld, C., de Kreij, C., van der Wees, A., 1991. Normen voor waterqualiteit in de glastuinbouw, No. 11, fifth
ed., p. 23.
Stanghellini, M.E., Rassmussen, S.L., 1994. Hydroponics: a solution for zoosporic pathogens. Plant Dis. 78,
1129–1138.
Stolk, A.P., 2001. Landelijk Meetnet Regenwatersamenstelling. Meetresultaten 2000. RIVM Rapport 723101 057/
2001, http://www.rivm.nl/bibliotheek/rapporten/723101057.html.
Tognoni, F., Pardossi, A., Serra, G., 1998. Water pollution and the greenhouse environmental costs. Acta Hort.
458, 385–394.
Van Assche, C., Vangheel, M., 1994. Special phytopathological problems in soilless cultures and substrate
cultures. Acta Hort. 361, 355–360.
Van Peer, R., Schippers, B., 1989. Plant growth responses to bacterization with selected Pseudomonas spp. strains
and rhizosphere microbial development in hydroponic cultures. Can. J. Microbiol. 35, 456–463.
Van Staden, J., 1999. Occurence and potential physiological effects of algal plant growth regulators. In:
Proceedings of the International Workshop and Training Course on Microalgal Biology and Biotechnology.
Mosonmagyarovar, Hungary, p. 40.
Vanachter, A., van Wambeke, E., van Assche, C., 1983. Potential danger for infection and spread of tomatoes in
hydroponics. Acta Hort. 133, 119–128.
Waechter-Kristensen, B., Gertsson, U.E., Sundin, P., 1994. Prospect for microbial stabilization in the hydroponic
culture of tomato using circulating nutrient solution. Acta Hort. 361, 382–387.
Wetzel, R.G., 2001. Limnology. Lake and River Ecosystems, third ed. Academic Press, San Diego.
Worch, E., 1997. Wasser und Wasserinhaltsstoffe. Eine Einführung in die Hydrochemie. B.G. Teubner Verlags-
gesellschaft, Stuttgart, Leipzig, p. 205.