ARTICLE IN PRESS

JOURNAL OF
FOOD COMPOSITION
AND ANALYSIS
Journal of Food Composition and Analysis 18 (2005) 635–645
www.elsevier.com/locate/jfca

Original Article

Flavonol content and composition of spring onions grown

hydroponically or in potting soil
L. Thompsona,, J. Morrisa, E. Peffleyb, C. Greenb, P. Paréc, D. Tissued,
R. Jasonic, J. Hutsonb, B. Wehnerd, C. Kaneb
a
Animal and Food Sciences, Texas Tech University, P.O. Box 42141, Lubbock, TX, 79409, USA
b
Plant and Soil Sciences, Texas Tech University, P.O. Box 42122, Lubbock, TX, USA
c
Chemistry and Biochemistry, Texas Tech University, P.O. Box 41061, Lubbock, TX, USA
d
Biological Sciences, Texas Tech University, P.O. Box 43131, Lubbock, TX, USA
Received 30 September 2003; received in revised form 5 March 2004; accepted 16 June 2004

Abstract

Two experiments were conducted as part of an effort to evaluate the suitability of onions as a candidate
crop for testing in a closed, controlled environment, hydroponic-based plant facility designed for long-term
manned space missions (NASA Engineering Development Unit). Composition and total flavonol content
of the plants were determined as they matured in a hydroponic-versus a soil-based system. ‘Purplette’
onions (Allium cepa L.) were grown hydroponically in a greenhouse for as long as 77 days. Composition of
the plant tissue was determined at weekly or biweekly intervals. Ca, Mg, K, and N (wet matter basis) all
decreased as plants matured. Dry matter (DM) and S contents were constant regardless of age averaging
10.6% and 187 mg/100 g, respectively. Total flavonol (TF) content increased as plants matured
(226–538 mg/100 g at 14 and 77 days, respectively). Onions grown in hydroponic units or in potting
medium had similar composition for all constituents examined (10.38%, 0.550%, 4.15%, and 0.97% DM,
N, C, and ash, respectively; and 126.0, 55.5, 270, 185 and 453 mg/100 g Ca, Mg, K, S and TF, respectively).
Based on phenotypic characteristics and composition determined in this study, onions were well suited to
hydroponic propagation.
r 2004 Elsevier Inc. All rights reserved.

Keywords: Allium cepa; Phytochemicals; Mineral composition; Hydroponic

Corresponding author. Tel.: +1-806-742-4862; fax: +1-806-742-4003.

E-mail address: leslie.thompson@ttu.edu (L. Thompson).

0889-1575/$ - see front matter r 2004 Elsevier Inc. All rights reserved.
doi:10.1016/j.jfca.2004.06.009
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636 L. Thompson et al. / Journal of Food Composition and Analysis 18 (2005) 635–645

1. Introduction

Food is an important nutritional, psychological and health aspect of any space mission.
Perchonok and Bourland (2002) report that as space missions become longer, the need for better
nutrition, increased food variety and ease of consumption becomes increasingly important. The
goal of food system development for long-duration missions is to provide acceptable foods that
taste similar to foods grown on earth while providing appropriate nutrition, and concurrently
minimizing food volume, mass and waste (Perchonok and Bourland, 2002). Not only there are
unique challenges in growing crops during missions and processing crops into foodstuffs, but also
many physiological effects occur in humans during long-term missions, which can impact food
system needs. During extended missions, physiological effects observed include weight loss, shift
in body fluids resulting in altered or diminished sensory acuity, dehydration, constipation,
electrolyte imbalance, Ca and K loss, decreased red blood cell mass, and motion sickness. These
physiological changes profoundly affect the acceptability of foods and the nutritional needs of
astronauts (Perchonok and Bourland, 2002).
Research efforts at the Texas Tech University Center for Space Science are underway to
provide NASA Advanced Life Science (ALS) program with horticultural strategies and cultivar
screening for onions and related Allium spp. (Peffley et al., 2003) for the controlled ecological life-
support system (CELSS), which utilizes closed growth systems for crop production to provide
support systems for humans in space (Kliss et al., 1993). Hydroponic propagation is one
production basis being utilized for CELSS units. These closed growth systems create growth
conditions quite different from those encountered in the field (Kliss et al., 1996). Among the
factors that vary are solid-support matrix, nutrient delivery and availability, relative humidity,
photoperiod, accumulation of volatiles, and CO2 partial pressure (Kliss et al., 1996). Such
differing conditions have been associated with variability in composition compared to field grown
crops (Alamazan et al., 1997; McKeehen et al., 1996; Steinberg et al., 2000; Wheeler, Mackowiak,
et al., 1993, 1994 & 1996). Some of the constituents that have been altered in wheat, soybeans,
lettuce, potatoes or sweet potatoes by hydroponic propagation include ash, protein, carbohydrate,
Ca, Mg, K, Na, S, and crude fiber (Alamazan et al., 1997; McKeehen et al., 1996; Steinberg et al.,
2000; Wheeler et al., 1993, 1994, 1996). Most of the information available on the nutritional
composition of edible plant tissues has been derived from field-grown crops and as such may not
be a reliable indication of the nutritional value of crops grown hydroponically.
Recent interest in the health benefits of phytochemicals has resulted in the examination of
flavonols, a group of secondary plant metabolites, flavonoids, which are non-nutritive plant
polyphenols widely reported as providing health benefits, mainly through antioxidant or anti-
inflammatory properties (Hertog, et al., 1992; Leighton et al., 1992; Greenfield, et al., 1993;
Rampton and Collins, 1993). The most abundant phytochemical in onions, quercetin, belongs to
the class of flavonoids called flavonols. A recently reported benefit of dietary flavonoids indicates
that these compounds may inhibit bone resorption (Wattel et al., 2003), which could be of benefit
in long-term missions under microgravity since a commonly reported problem associated with
long-term missions is the loss of Ca and decreased bone density (Perchonok and Bourland, 2002).
Some onion cultivars have been reported to contain abundant levels of flavonols and might be
especially important for extended missions. Limited studies have been conducted on the
phytochemical aspects of hydroponically grown crops, including onions.
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The objectives of the present study were to determine the effects of plant age at harvest and
hydroponic growth, on the total flavonol (TF) content and composition of spring onions. The
specific experimental objectives were to determine the effect of: (1) plant age (Experiment 1); and
(2) growth media (hydroponic vs. potting soil) and plant age (Experiment 2), on the phenotypic
variables (edible biomass, plant length, and number of leaves), TF content and composition of
spring onions.

2. Materials and methods

2.1. Plant materials

2.1.1. Experiment 1
Two-hundred long-day onions (Allium cepa L. ‘Purplette’) (Johnny’s Selected Seeds; Albion,
ME, USA) were planted in Oasis Hortcubest (Smithers-Oasis, Kent, OH, USA) at a depth of
0.75 mm. Plants were germinated and grown in the Texas Tech University Horticulture
greenhouse between March and July 2001, and irrigated daily with Hydro-Solt (Scotts-Sierra
Horticultural Product Co, Marysville, OH, USA). Twenty plants were harvested at 14, 21, 28, 35,
42, 49, 63, and 77 days after germination. The experiment was replicated seven times with
plantings occurring at weekly intervals.

2.1.2. Experiment 2
Four-hundred long-day onions (A. cepa ‘Purplette’) (Johnny’s Selected Seeds; Albion, ME,
USA) were planted in Oasis Hortcubest at a depth of 0.75 mm (Smithers-Oasis, Kent, OH, USA)
to create two replications of 200 plants per replication within the hydroponic treatment. Another
400 long-day onions (A. cepa ‘Purplette’) (Johnny’s Selected Seeds; Albion, ME, USA) were
planted in Ball Growing Mix 2t (Ball Seed; West Chicago, IL, USA) potting soil at a depth of
0.75 mm with row and seed spacing matching that of the Oasist (3.8 cm
3.8 cm) to create two
replications of 200 plants per replication within the potting soil treatment. Plants were germinated
and grown in the Texas Tech University Horticulture greenhouse between May and August 2001.
After germination, plants were irrigated daily with Hydro-Solt (Scotts-Sierra Horticultural
Product Co, Marysville, OH, USA). Forty plants (20 plants per replication) from each growing
medium were harvested weekly at 14, 21, 28, 35, 42, and 49 days after germination.

2.2. Phenotypic measurements and sample preparation

At harvest, roots were removed at the basal plate and height, edible biomass (weight) and net
number of leaves were determined on the remaining portion (leaves and bulb). In order to have
sufficient material for analytical purposes, plants were composited depending on plant mass. Ten
plants harvested at 14 and 21 days were composited to make two samples representative of the
respective age. Five plants were composited at 28 d and 35 d, four plants were used at 42 and 49
days, and two plants were used at 63 days of age. Plants at 77 days were large enough to constitute
a single sample. After weighing, plants were chopped into pieces p5 cm and frozen in liquid
nitrogen. The frozen plant material was ground to a fine powdery consistency in a coffee grinder
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638 L. Thompson et al. / Journal of Food Composition and Analysis 18 (2005) 635–645

(Brauns; Boston, MA, USA; Mr. Coffees; Hattiesburg, MS, USA). Approximately a 2-g sample
of ground onion was used immediately for flavonol extraction and the remaining powder was
placed in Whirlpackt bags for storage in a –20 1C-freezer for subsequent analysis.

2.3. Total flavonol determination

Flavonols were extracted using 80% ethanol (Lombard, et al., 2003). Filtrate was collected in 2-
mL Eppendorf tubes and stored in a –20 1C freezer. Immediately prior to analysis, extracts were
warmed to room temperature, and a 0.5-mL sample was diluted with 4.5 mL of 80% ethanol.
Absorbance of the diluted solution was determined on a Bausch & Lomb Spectronic 20
Colorimeter/Spectrophotometer Model 95 (Richmond, BC, Canada) at 362 nm in 1-cm cuvettes
(Fisher Scientific; Houston, TX, USA; Bausch & Lomb; Richmond, BC, Canada) (Lombard et
al., 2003). Duplicate readings were preformed on each sample and averaged.
Isoquercitrin (Extrasynthese; Genay, France) was used as a standard (Lombard, 2000) and nine
different standards with concentrations ranging from 0.00625 to 0.125 mg/mL were used to create
a standard curve (absorbance vs. concentration) using linear regression. Total flavonol content
was calculated on a wet matter basis using the equation for the best fitting line.

2.4. Dry matter

Dry matter content was determined by drying duplicate samples in a vacuum oven (National
Appliance Co.; Portland, OR, USA) 16 h at 100 1C (AOAC, 1995).

2.5. Nitrogen and carbon analysis

Total N and C percentage were determined in duplicate on 2–4 mg dried samples with a Carlo
Erba NCS 2500 C/N analyzer (Milan, Italy) and reported on a wet matter basis. Atropine (CE
Elantech; Lakewood, NJ, USA) was used as standard.

2.6. Ash and mineral content

Duplicate samples were dried in a vacuum oven as previously described and incinerated in a
muffle furnace (Sybran Thermolyne 2000 Furnace, Thermolyne Corp.; Dubuque, IA, USA; Blue
M Lab-Heat Muffle Furnace, Blue M Electric Co.; Blue Island, IL, USA) for 16 h at 500 1C
(AOAC, 1995). Calcium, Mg and K contents were determined on the ash residues using atomic
absorption spectroscopy (Perkin, 1976; AOAC, 1995). Ash residues were dissolved in 15 mL of
20% HNO3 for at least 5 h. Each solution was filtered through Whatman 40 (Clifton, NJ, USA)
ashless filter paper and diluted to a total volume of 100 mL with D-H2O in a volumetric flask. One
mL aliquots of each sample were placed in two separate tubes. To one set of tubes 10 mL of
distilled water was added. To the other set 9.5 mL of D-H2O and 0.5 mL of 5% lanthanum
chloride were added as releasing agent for use in Ca and Mg quantification. Concentrations of the
minerals were determined using a Perkin Elmer 2380 atomic absorption spectrophotometer
(Norwalk, CT, USA). Calcium, Mg or Na–K Intensitron hollow cathode lamps were used (Perkin
Elmer, Norwalk, CT, USA) at wavelengths of 422.7, 285.2, or 766.5 nm, respectively, with
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L. Thompson et al. / Journal of Food Composition and Analysis 18 (2005) 635–645 639

oxidizing lean, blue flame air/acetylene flames. Ash and minerals were reported on a wet matter
basis.

2.7. Sulfur quantification

Sulfur content was determined using an AOAC (1995) oxidative gravimetric method consisting
of the 923.01 magnesium nitrate and the 920.10 sodium peroxide methods. Approximately 1 g of
fresh ground onion was weighed in a ceramic crucible. At the completion of the oxidation the
remaining residue (BaSO4) was weighed and multiplied by 0.1374 to determine total sulfur content
present in the sample. The S content (as a percentage of wet matter) was reported.

2.8. Experimental design and statistical analysis

2.8.1. Experiment 1
Data were analyzed as an age at harvest (6)
planting date (Seven replications) factorial for all
variables with the exception of TF, which was analyzed as an 8
7 factorial. Analysis of variance
was conducted using general linear models (Proc GLM) in SAS 8.0 (SAS Institute, Cary, NC,
USA). The height, weight, and leaf number means for the edible biomass 7 standard error of the
means (S.E.M.) were presented. No significant interaction between age at harvest and replication
was found for the composition attributes; therefore, age of plant variables means were separated
using Tukey’s mean separation (Po 0.05).

2.8.2. Experiment 2
Data were analyzed as a 6 (age at harvest)
2 (growth media) factorial with age of harvest at
14, 21, 28, 35, 42 and 49 days, and two growing media, Oasis Hortcubest and Ball Growing Mix
2t potting soil. Analysis of variance was conducted using general linear models (Proc GLM) in
SAS 8.0 (SAS Institute, Cary, NC, USA). The height, weight, and leaf number means for the
edible biomass7standard error of the means (S.E.M.) were presented. No significant interaction
between age at harvest and growth media were found for any of the variables. As such main effect
means were separated using Tukey’s mean separation (Po0.05).

3. Results

3.1. Experiment 1—plant age

Age at harvest did not significantly affect DM or S contents of ‘Purplette’ onions, averaging
10.6 %70.10, and 18773.65 mg/100 g, respectively (Table 1). Percentage N significantly
decreased as age at harvest increased (Table 1). Percentage C decreased during the time period
of from 28 to 77 days of age (Table 1). Percentage ash significantly increased as plants aged
however; Ca, Mg and K levels decreased from early high concentrations at 28 days of age to their
lowest values at 77 days of age (Table 1). A sharp, significant decrease in concentration for Ca,
Mg, and K occurred from 49 to 63 days of age. Total flavonol content significantly increased
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640 L. Thompson et al. / Journal of Food Composition and Analysis 18 (2005) 635–645

Table 1
Total flavonol (TF) content and composition of edible portions (expressed on a wet matter basis) of ‘Purplette’ onions
(A. cepa) grown hydroponically in a greenhouse from March to July 2001a

Age (days)

Variable n 28 35 42 49 63 77

Dry matter (%) 266 10.6a 10.5a 10.7a 10.6a 10.6a 10.5a
Nitrogen (%) 253 0.55a 0.55a 0.53b 0.51b 0.46c 0.41d
Carbon (%) 266 4.16a,b 4.20a,b 4.12b 4.11b 4.10b 4.00c
Ash (%) 253 0.91c 0.95c,d,e 0.92d,e 0.98b,c,d 1.02a,b 1.07a
Ca (mg/100 g) 253 124a 125a 123a 129a 85.7b 65.6c
Mg (mg/100 g) 253 56.8a 57.1a 54.1b 53.8b 24.0c 22.3d
K (mg/100 g) 253 274a 270a 266a 265a 229b 223b
S (mg/100 g) 201 185a 187a 188a 189a 188a 185a
a
Means within a row with different letters are different (Po0.05).

600.0
538.2 a
512.1 a 529.0 a
480.4 b
500.0 457.2 b
414.4 c
TF (mg /100 g)

400.0
334.5 d

300.0
226.1 e
200.0

100.0

0.0
14 21 28 35 42 49 63 77
Age (days)

Fig. 1. Total flavonol (TF) content in edible portions of ‘Purplette’ onions (A. cepa) grown hydroponically in a
greenhouse from March to July 2001 ðn ¼ 88Þ. Means with different letters differ ðPo0:05Þ.

throughout the growing period, increasing about 145% during the 77-day experimental period,
reaching a high of 538 mg/100 g (Fig. 1).
Leaf number doubled during the period from 14 to 77 days, and the height of the plants
increased nearly five-fold during this same period (Table 2). In the initial growth period up to 21
days edible plant mass changed little, but in the 1-week period between day 21 and 28, edible
biomass increased 800% and plant height increased by 12.41 cm (Table 2). During a 5 week-
period between day 28 and 63, plant height doubled and mass increased five-fold (Table 2).
Growth rate slowed between 63 and 77 days (Table 2).
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Table 2
Phenotypic characteristics of edible portion (leaves and bulb) of ‘Purplette’ onions (A. cepa) grown hydroponically in a
greenhouse from March to July 2001

Age (day) 14 21 28 35 42 49 63 77

N 140 140 140 140 140 140 120 80

Leaf number 2.00 2.00 2.56 3.40 3.61 3.20 4.14 4.39
S.E.M. 0.00 0.00 0.05 0.05 0.06 0.06 0.07 0.90
Height (cm) 7.87 6.84 19.25 25.75 30.14 33.37 38.30 39.09
S.E.M. 0.02 0.03 0.33 0.23 0.36 0.36 0.35 0.51
Weight (g) 0.07 0.09 0.81 1.07 2.01 2.76 4.19 4.54
S.E.M. 0.00 0.00 0.02 0.02 0.04 0.05 0.08 0.09

Table 3
Total flavonol (TF) content and composition of edible portions of ‘Purplette’ onions (A. cepa L.) grown hydroponically
(H) and in potting soil (PS) in a greenhouse from May to August 2001a

Treatment DM(%) N(%) C(%) Ash(%) Ca(mg/100 g) Mg(mg/100 g) K(mg/100 g) S(mg/100 g) TF(mg/100 g)
n 72 72 72 72 69 69 69 66 88

Age (day)
14 — — — — — — — — 262c
21 — — — — — — — — 343c
28 9.78a 0.575a 4.16a 0.92b 128a 56.9a 272a 183a 463b
35 10.5a 0.560a,b 4.16a 0.97a 124a 57.1a 270a 186a 528a,b
42 10.7a 0.544b 4.15a 1.00a 126a 54.1a 269a 181a 538a
49 10.5a 0.526c 4.12a 1.00a 126a 53.9a 268a 190a 583a
Growth media
H 10.2a 0.552a 4.16a 1.00a 126a 55.2a 272a 186a 452a
PS 10.5a 0.547a 4.14a 0.94a 126a 55.8a 268a 184a 454a
a
Means within a variable and main effect with different letters are different (Po0.05). Insufficient plant material was
available for analyses for day 14 and 21 with the exception of TF.

3.2. Experiment 2—hydroponic vs. potting soil

No significant interaction was found between age at harvest and growth media for any of the
variables, thus means of main effects are in Table 3. Propagation in hydroponics vs. potting soil
had no effect on any of the variables (Table 2). Up to 49 days of age, means for DM and C
percentage, Ca, Mg, K, and S content were similar regardless of plant age (Table 2) and averaged
10.4%, 4.15%, 126, 55.5, 270, and 185 mg/100 g, respectively. Percentage N significantly
decreased with plant age (Table 2). Between 28 and 35 days ash content increased but then
remained constant. Total flavonol content significantly increased by 122% over the 35-day
experimental period.
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4. Discussion

The USDA nutrient database indicates that raw spring onions (A. cepa or fistulosum) have the
following composition: DM=10.17%, N=0.29%, ash=0.81%, Ca=72 mg/100 g, Mg=20 mg/
100 g, and K=276 mg/100 g (US Department of Agriculture,Agricultural Research Service,
USDA, 2002). The average DM content reported in the present study was slightly higher than that
found in the US Department of Agriculture,Agricultural Research Service USDA (2002) nutrient
database. The N content regardless of age was higher than the USDA reported value. The
percentage ash in the present study was also slightly higher than the USDA reported values
especially after 49 days of age. Ca and Mg values reported in the USDA database were similar to
values at 63 to 77 days of harvest. Discrepancies between data in the present experiments and the
USDA database would be expected and could be a result of a wide variety of factors such as plant
age, cultivar, environmental conditions, nutrient availability, and storage conditions of the
harvested plants.
In Experiment 1 during the 49–63 days sampling period, the onions appeared to start the
bulbing process (observed by the investigators but not specifically measured), which coincided
with significant changes that occurred in percentage N, ash, Ca, Mg and K. The biochemical
changes during bulbing are not completely understood but seem to influence composition. Randle
(2000) and Brewster (1994) have previously reported N content declines as plants age and the
bulbing process is initiated, while C content tends to increase as a result of accumulation and
polymerization of non-structural carbohydrates during bulbing (Kahane et al., 2001). Brewster
(1994) has hypothesized that increases in reducing sugars, sucrose, and fructans during bulbing
can lead to changes in the osmotic potential which could then affect the mineral content
(Brewster, 1994).
In mature, cured bulb onions, Lombard (2000) reported TF levels in five cultivars ranging from
28.5 to 58.0 mg/100 g, at least 10-fold less than the values found in the present study. Other studies
have reported quercetin levels in mature bulb onions ranging from 137 to 178 mg/100 g (Price and
Rhodes, 1997), 5.4 to 28.6 mg/100 g (Patil, et al., 1995), while Hertog and coworkers (1992)
reported a mean quercetin content of 34.7 mg/100 g. Lombard (2000) estimated that about 90% of
the TF in bulb onions are quercetin conjugates. The observed TF content in the present study
compared to the TF cited from previous studies could be related to the fact that the onions in this
study were spring onions rather than cured bulb onions and that both the leaf and bulb parts were
analyzed in this study.
Constituent concentrations that have been previously reported to be different between soil-
grown and hydroponically grown plants include ash, protein, carbohydrate, Ca, Mg, K, Na, S,
and crude fiber (Alamazan et al., 1997; McKeehen et al., 1996; Steinberg et al., 2000; Wheeler et
al., 1993, 1994, 1996). These differences have been attributed to different nutrient content and
availability between growth media (Taiz and Zeiger, 1998, Resh 1991). Resh (1991) has suggested
that differences in ash and mineral content between soil-grown and hydroponically grown plants
may be related to ion availability. Soil particles may attract and bind cations reducing the
availability of the ions for use by the plants. Oasis, the solid support often used in hydroponics, is
chemically inert (Resh, 1991) and does not interfere with mineral absorption, thereby allowing
plants to have unlimited uptake of minerals, and increase the overall ash content within the plant
tissues.
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Table 4
Phenotypic characteristics of edible portions (leaves and bulb) of ‘Purplette’ onions (A. cepa) grown hydroponically (H)
and in potting soil (PS) in a greenhouse from May to August 2001

Growth media Age (day) 14 21 28 35 42 49

H n 40 40 40 40 40 40

Leaf number 2.00 2.00 3.05 3.93 4.08 3.08

S.E.M. 0.00 0.00 0.12 0.09 0.10 0.12
Height (cm) 7.54 9.67 23.93 26.50 28.46 25.72
S.E.M. 0.06 0.05 0.68 0.40 0.49 0.56
Weight (g) 0.07 0.09 0.99 1.20 1.85 1.43
S.E.M. 0.00 0.00 0.08 0.04 0.05 0.08
PS
Leaf number 2.00 2.00 3.80 4.05 4.25 3.93
S.E.M. 0.00 0.00 0.13 0.11 0.11 0.12
Height (cm) 7.74 9.65 24.46 27.94 29.68 26.53
S.E.M. 0.05 0.08 0.60 0.41 0.55 0.59
Weight (g) 0.08 0.09 1.39 1.67 2.01 2.31
S.E.M. 0.00 0.00 0.09 0.05 0.06 0.16

In the present study, no differences were found in composition between plants grown
hydroponically and in potting soil (Table 4). One factor that may account for a lack of differences
is that potting soil-grown plants were watered with the same nutrient solution, Peter’s Hydro-Sol,
as was used as the nutrient solution for the hydroponic units. Another factor that may have lead
to no significant differences between treatments is that the hydroponic system used in the present
study was a static system. In the majority of other studies, hydroponically grown plants were
grown in a dynamic flow systems, in which the nutrient solutions were circulated around the root
mass continuously. Because all plants in the present study were exposed to the same nutrient
solution (but different support matrix) it is not unexpected that the plant composition was similar,
even though the solid support matrix was different.

5. Conclusions

Plant age at harvest can affect composition and needs to be considered when managing
nutritional needs of astronauts or when compiling composition data for nutrient databases. Key
constituents in spring onions that varied by plant age were N, C, ash, Ca, Mg, K and TF.
Hydroponically grown spring onions under the conditions in this study differed little in
composition from potting soil-grown counterparts. Further studies need to examine the plant
composition as affected by static and dynamic hydroponic systems and by different nutrient
solutions.
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Acknowledgements

Support for this research was provided by a grant from the National Aeronautics and Space
Administration, cooperative agreement JSC-4-01-9729.

References

AOAC, 1995. Official Methods of Analysis, 16th ed. AOAC International, Arlington, VA.
Alamazan, A.M., Begum, F., Johnson, C., 1997. Nutritional quality of sweet potato greens from greenhouse plants.
Journal of Food Composition and Analysis 10, 246–253.
Brewster, J.L., 1994. Onions and Other Vegetable Alliums. University Press, Cambridge.
Greenfield, R.J., Pernchard, N.A., Teare, J.P., Thompson, R.P., 1993. The mode of action of the aminosalicylates in
inflammatory bowel disease: a review. Alimentary Pharmacology and Therapeutics 7, 321–369.
Hertog, M.G.L.S, Hollman, P.C.H., Katan, M.B., 1992. Content of potentially anticarcinogenic flavonoids of 28
vegetables and 9 fruits commonly consumed in the Netherlands. Journal of Agricultural and Food Chemistry 40
(12), 2379–2383.
Kahane, R., Vialle-Guerin, E., Boukema, I., Tzanoudakis, D., Bellamy, C., Chamaux, C., Kik, C., 2001. Changes in
non-structural carbohydrate composition during bulbing in sweet and high-solid onions in field experiments.
Environmental and Experimental Botany 45, 73–83.
Kliss, M., Borchers, B. & Drews, M. (1993). Design considerations for the CELSS test facility engineering development
unit. SAE Technical Paper Series 932124, 23rd International Conference on Environmental Systems. Colorado
Springs, CO, July 12–15.
Kliss, M., Blackwell, C., Zografos, A., Drews, M., MacElroy, R., McKenna, R., Heyenga, G., 1996. Initial closed
operation of the CELSS test facility engineering development unit. Advanced Space Research 31 (1), 263–271.
Leighton, T., Gaither, C., Fluss, L., Harter, W.K., Canasado, J., Notario, V., 1992. Molecular characterization of
quercetin and quercetin glycosides in Allium vegetables. American Chemical Society 16, 221–238.
Lombard, K.A., 2000. Investigation of the flavonol quercetin in onion (Allium cepa L.) by high-performance liquid
chromatography (HPLC) and spectrophotometric methodology M.Sc. Thesis, Texas Tech University, Lubbock,
TX, 41–42p; Available from: University Library, Lubbock TX; AC805.T3 2000, no. 147.
Lombard, K., Geoffriau, E., Peffley, E., 2003. Quantification of quercetin in onion by HPLC and spectrophotometric
analyses. HortScience 37 (4), 682–685.
McKeehen, J.D., Smart, D.J., Mackowiak, C.L., Wheeler, R.M., Nielsen, S.S., 1996. Effects of CO2 levels on nutrient
content of lettuce and radish. Advanced Space Research 18 (4/5), 85–92.
Patil, B.S., Pike, L.M., Yoo, K.S., 1995. Variation in the quercetin content in different colored onions (Allium cepa L.).
Journal of American Society Horticultural Science, 120, 909–913.
Peffley, E., Green, C., Pare, P., Thompson, L., Tissue, D., 2003 Plant carbon allocation and partitioning of metabolites
in a closed growth facility. Bioastronautics Investigators’ Workshop. January 13–15, 2003, Galveston, TX; National
Aeronautics and Space Administration, Houston, TX, pp. 120.
Perchonok, M., Bourland, C., 2002. NASA food systems: past, present and future. Nutrition 18, 913–920.
Perkin E., 1976. Analytical Methods for Atomic Absorption Spectrophotometry. Perkin Elmer, Norwalk, CT.
Price, K.R., Rhodes, M.J.C., 1997. Analysis of the major flavonol glycosides present in four varieties of onion (Allium
cepa) and changes in composition resulting from autolysis. Journal of Science and Food Agriculture 74, 331–339.
Rampton, D.S., Collins, C.E., 1993. Thromboxanes in inflammatory bowel disease: pathogenic and therapeutic
implications. Aliment Pharmacological Therapy 7, 357–367.
Randle, W.M., 2000. Increasing nitrogen concentration in hydroponic solutions affects onion flavor and bulb quality.
Journal of American Society of Horticultural Science 125 (2), 254–259.
Resh, H.M., 1991. Hydroponic food production, 3rd ed.. Woodbridge Press Publishing Co., Santa Barbara 1–65.
Steinberg, S.L., Ming, D.W., Henderson, K.E., Carrier, C., Gruener, J.E., Barta, D.J., Henninger, D.L., 2000. Wheat
responses to differences in water and nutritional status between zeoponic and hydroponic growth systems.
Agronomy Journal 92, 353–360.
 ARTICLE IN PRESS

L. Thompson et al. / Journal of Food Composition and Analysis 18 (2005) 635–645 645

Taiz, L., Zeiger, E.I., 1998. Plant defenses: surface protectants and secondary metabolites. In: Taiz, L., Zeiger, E.I.
(Eds.), Plant physiology, 2nd ed.. Sinauer Associates, Inc., Sunderland, pp. 342–376.
US Department of Agriculture, Agricultural Research Service, USDA, 2002. USDA National Nutrient Database for
Standard Reference, Release 15. Nutrient Data Laboratory Home Page, http://www.nal.usda.gov/fnic/foodcomp.
Wattel, A., Kamel, S., Mentaverri, R., Lorget, F., Prouillet, C., Petit, J.P., Fardelonne, P., Brazier, M., 2003. Potential
inhibitory effect of naturally occurring flavonoids quercetin and kaempferol on in vitro osteoclast bone resorption.
Biochemical Pharmacology 65 (1), 35–42.
Wheeler, R.M., Mackowiak, C.L., Sager, J.C., Knott, W.M., Berry, W.L., 1994. Proximate nutritional composition of
CELSS crops grown at different CO2 partial pressures. Advance Space Research 14 (11), 171–176.
Wheeler, R.M., Mackowiak, C.L., Sager, J.C., Knott, W.M., Berry, W.L., 1996. Proximate nutritional composition of
CELSS crops grown in NASA’s biomass production chamber. Advance Space Research 18 (4/5), 43–47.
Wheeler, R.M., Mackowiak, C.L., Siegriest, L.M., Sager, J.C., 1993. Supraoptimal carbon dioxide effects on growth of
soybean [Glycine max (L.) Merr.]. Journal of Plant Physiology 42, 173–178.