Plant Physiol.
(1991) 95, 1125-1130
0032-0889/91/95/1125/06/$01 .00/0
Nonrecirculating Hydroponic System Suitable for Uptake
Studies at Very Low Nutrient Concentrations'
Vincent P. Gutschick* and Lou Ellen Kay
Department of Biology, New Mexico State University, Las Cruces, New Mexico 88003
ABSTRACT
We describe the mechanical, electronic, hydraulic, and struc-
tural design of a nonrecirculating hydroponic system. The system
is particularly suited to studies at very low nutrient concentra-
tions, for which on-line concentration monitoring methods either
do not exist or are costly and limited to monitoring relatively few
individual plants. Solutions are mixed automatically to chosen
concentrations, which can be set differently for every pump fed
from a master supply of deionized water and nutrient concen-
trates. Pumping rates can be varied over a 50-fold range, up to
400 liters per day, which suffices to maintain a number of large,
post-seedling plants in rapid growth at (sub)micromolar levels of
N and P. The outflow of each pump is divided among as many as
12 separate root chambers. In each chamber one may monitor
uptake by individual plant roots or segments thereof, by meas-
uring nutrient depletion in batch samples of solution. The system
is constructed from nontoxic materials that do not adsorb nutrient
ions; no transient shifts of nitrate and phosphate concentrations
are observable at the submicromolar level. Nonrecirculation of
solutions limits problems of pH shifts, microbial contamination,
and cumulative imbalances in unmonitored nutrients. We note
several disadvantages, principally related to high consumption of
deionized water and solutes. The reciprocating pumps can be
constructed inexpensively, particularly by the researcher. We
also report previously unattainable control of passive temperature
rise of chambers exposed to full sunlight, by use of white epoxy
paint.
Long-term solution culture at micromolar levels of nu-
trients is required in studies of adaptations of nutrient uptake
systems and root growth under a variety of stresses. Full
expression of growth adaptations, such as in root:shoot ratio,
requires growth to considerable size (9). Large, vigorously
growing plants will deplete large volumes of nutrient solution.
We take an example from our recent experiments and con-
sider a plant having a dry mass m = 2 g, a tissue phosphorus
content of 0.2% by dry weight (fractional contentf= 0.002),
and a relative growth rate R of 0.15 d-'. Consider further
providing phosphorus (atomic weight M = 31 g molV') as
phosphate at a concentration of 1 mmol m-3. If one allows a
30% relative concentration depletion, Ac = 0.3 mmol m-3
(0.3 uM), the required flow rate J is determined by solving the
equation between plant uptake and solution depletion:
Mass of P taken up per unit time = mfR JAcM. (1)
= exposed to full sun to remain within 1°C of ambient air
'Funding by the U. S. Department of Energy, Ecological Research.
1125
Received for publication August 27, 1990
Accepted December 11, 1990
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With the given parameter choices, the required flow rate is
65 L/d for every replicate plant. Most hydroponic systems
capable of providing such large volumes do so by recirculating
solution from a large reservoir (2, 3), often with automated
makeup of depleted nutrients to maintain concentrations and
to measure uptake (e.g. 7, 13, 14). Recirculation can increase
problems of microbial contamination. It also requires auto-
mated controls for makeup of nutrient ions or of hydrogen
ions (pH balance) during nitrate or ammonium usage. Con-
centration sensors for the controls are commonly based upon
ion-selective electrodes (4, 7) or, rarely, flow-injection analysis
(5). Electrodes are adequately sensitive for nitrate down to 5
,AM (4) but not down to (sub)micromolar levels at which one
may wish to maintain other nutrients, such as phosphate and
the micronutrients. Flow-injection systems are quite costly
and can monitor relatively few individual plants or groups at
a time (for example, 8 plants, in the case of ref. 5, compared
with the 28 to 84 plants we survey with our system). Typically,
recirculating systems monitor relatively few nutrients and fail
to monitor most of the nutrients, particularly micronutrients.
The requirement to oversupply the unmonitored nutrients
might cause initial toxicity problems in multiweek experi-
ments with nutrients such as copper.
We have designed our system to mix, automatically on
demand, large volumes of fresh solution for single-pass flow.
The contents of all nutrients at inputs to every plant are
controlled by mixing ratios between three solutions. The
average concentrations at the plant root system are drawn
down from input levels by nutrient uptake, but adequate
pumping rates can hold all concentrations within relatively
tight limits. The design addresses several other major practical
problems, foremost that of providing inexpensive, well-regu-
lated pumping for multiple treatments and replicates. (The
pumps may also be used to advantage in recirculating sys-
tems.) We have designed and built positive-displacement
pumps based on 140-mL plastic syringes, with electronically
controlled repetition rates. Simple hydraulic resistance divid-
ers apportion flows reproducibly to replicate plants. Our
growth containers are divided into three separate chambers
meeting in the center, allowing two- and even three-way
divisions of root systems for differential treatment. We also
have identified a coating that is highly reflective at short
wavelengths and high in thermal emissivity, allowing surfaces
temperature. The entire system is constructed of nontoxic
materials (10) that are also low in capacity to release or adsorb
nutrient ions.
1126 GUTSCHICK AND KAY Plant Physiol. Vol. 95, 1991

This report extends and clarifies an earlier presentation

(1 1).

MATERIALS AND METHODS

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Figure 1 shows the full system schematically. Full mechan-
ical and electronic plans are available from us. optional 1
Nutrient solutions are mixed automatically in two stages. her
First, the system mixes a base solution of all the elements not
be varied in concentration between experimental groups, that
is, all elements but N and P in our case. The invariant
elements are provided as two 90x concentrates (A, B in Table bO
I), designed to be soluble separately. The A and B concentrates aE
are mixed on demand with DIW2 provided at nearly line
.2
pressure through two large mixed-bed deionizer tanks in series
(each with commercial designation of 10,500-grain capacity
[as CaCO3J, equivalent to 7 mol of divalent anions and 7 mol
divalent cations). Outflows of both deionizer tanks are mon-
itored for specific resistivity to give timely warning of need
for replacement. The mixed base solution is held in a 20-L
polyethylene carboy suspended on a metal spring. When the
carboy is depleted to 7 L, it rises to trip a microswitch. A
latching relay then activates (via solid-state relays) several
solenoid valves in order to: (a) empty the contents of two 150-
mL reservoirs of A and B base concentrates into the base-
solution carboy; it is important to maintain a 7-L reserve so
that incoming concentrates do not precipitate on contact; (b)
shut off the gravity-fed refilling of the 150-mL concentrate
reservoirs; (c) turn on the DIW inflow and activate a magnetic
stirrer mounted on the carboy bottom; and (d) shut off
outflow from the carboy, so that continued consumption of
base solution by pumps does not alter the concentration as
mixing occurs over the 3-min refill time. A second base
solution carboy, mounted lower, provides solution for pumps
while refilling of the first carboy proceeds. When the base
carboy refills to 20 L, it trips a second microswitch that
deactivates the solenoids noted above. It also trips a second
latching relay that opens inflow valves to refill the A and B
concentrate reservoirs, by gravity flow from 20-L carboys
mounted above. Inflow ceases when the liquid level in a
reservoir bridges two stainless steel pins, closing a low-current
relay circuit that shuts off solenoid valves.
The adjustable-concentration nutrients (e.g. N and P) are
held as 50-fold concentrates in 20-L carboys; they are mixed
with base solution at the pump. A reciprocating crank operates
three syringes. The large syringe (Monoject3 140-mL plastic
disposable; Sherwood Medical, DeLand, FL) is driven with a
12.5-cm stroke and delivers 95 mL of base solution per cycle.
The Leur-Lock fitting is bored out to reduce pumping back-
pressure, greatly prolonging syringe life to over 5000 cycles
before leakage becomes significant. Two glass syringes (Bec-
ton-Dickinson, 5-mL) operate on a 1.8-cm stroke and each Figure 1. Schematic diagram of hydroponic system. See text for full
delivers 2 mL of N or P concentrate per cycle. A single crank explanation. Solution flows along lines indicated in boldface. Dashed
arm holds the glass syringes in a rubber pad to eliminate lines indicate electrical connections. Solenoid-controlled valves are
noted as either normally open (N.O.) or normally closed (N.C.). System
2
Abbreviations: DIW, deionized water; FeEDTA, ferric ethylene- is shown configured for seven separate pumps, as used by authors.
diaminetetracetic acid; VAC, volts-alternating current.
3The mention of a specific produce name is not a unique endorse-
ment of this product. It does indicate satisfactory performance of the
specific product in long-term operation.
 HYDROPONIC SYSTEM
Table I. Composition of Nutrient Solutions
From concentrate "A," diluted 1:90
CaCI2,2.0 mM; Ca2, 2 mM; Cl, 4 mM
FeNaEDTA- 1.5 H20, 3.53 mg L-1; Fe3 , 0.5 Ag/ml (=9 MM)
From concentrate "B," diluted 1:90
K2SO4, 1.5 mM; K+, 3 mM; SO2-, 2.5 mM
MgSO4.7H20, 1 mM; Mg2+, 1 mM
MnSO4.7H20, 3.02 mg L-1; Mn2+, 0.6 ,g/ml (11 MM)
H3B03, 3.42 mg L-1; B, 0.6 Mg/ml (z55 MM)
ZnSO4-7H20, 1.32 mg L-1; Zn2 , 0.3 Mg/ml (-5 MM)
Na2MoO4.2H20, 0.126 mg L-1; Mo, 0.05 ,g/ml (-0.5 AM)
CuSO4.5H20, 0.078 mg L-'; Cu2+, 0.02 Mg/ml (=0.3 MM)
From concentrates for specific treatments, diluted 1:60
Treatment
1 2 3 4 7 6 5
Input N03 [Mm, from Ca(NO3)] 10 10 10 100 150 10 3
Input "PO4" [MM, from KH2PO4] 0.1 0.3 0.3 1 3 3 3
P-stress series - filter. All of the root chambers fed by a single pump must be
N-stress
series -.
breakage. Three pairs of stainless-steel check valves prevent
backflow at both entry and exit ports of each syringe. The
valves (NuPro 1/3-psi) are only lightly spring-loaded to keep
refilling drag low, and they have negligible void volume.
Valves must be checked individually every several days for
back-leakage; the Neoprene flap can be reseated. The full
crankshaft is bearing-mounted on both ends of its axis and is
machined from brass. The crank and motor mounts and the
base are machined from '/2-inch- (12.7 mm) thick stainless
steel for resistance to corrosion and stability against torques.
Four posts, one at each corner of the pump base, fit into other
pump bases, allowing stacking up to four pumps in one
footprint area.
The crankshaft is turned by a 3-rpm, high-torque, gear-
reduction motor (Jakel Motors model BG5P3K; Highland,
IL) operating at 1 7 VAC. Pumping rate is adjusted by setting
the interval between full cycles, to any value between 20 and
990 s in 10-s increments. This interval is controlled separately
for each pump by an integrated-circuit timer reading the
settings on a two-digit thumbwheel display. The timer may
control the entire revolution of a geared stepping motor
(Superior Electric, Bristol, CT) or, as in our system, it may
start an asynchronous motor. Revolution time of the latter
varies slightly with load, so that we use a crankshaft detent
and microswitch (details available) to provide power for an
exact 360° rotation, irrespective of load.
Outflows of the three syringes pumping base, N, and P
solutions merge in Tygon tubing linked with rigid Y-connec-
tors. The combined outflow entrains about 5 mL of air,
derived from (reproducible) air leakage past the large syringe's
piston during filling and perhaps from cavitation. Because air
bubbles can randomly plug the capillary-tube flow dividers
(described shortly), the air is removed by a simple standpipe
(of dark amber glass, to minimize chances of algal growth).
The flow from any single pump is then divided between 4
to 12 separate root chambers, representing individual plants
1127
or distinct sections of their root systems. Inflow to each
chamber goes through a uniform hydraulic resistance pro-
vided by a 10-cm length of 1-mm-diameter glass capillary
tubing. Only rapid flows such as provided by an intermittent
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reciprocating pump can be hydraulically divided accurately.
Without flow division, a separate positive-displacement pump
would be needed for each plant, at prohibitive cost or com-
plexity of operation.
One 2-L polystyrene beaker is divided into three root
chambers (Fig. 2) by acrylic plastic pieces held in place with
silicone sealant. The beaker lid of 1/2-in- (1 2.7-mm-)thick
acrylic plastic is cut to hold either (a) one plant positioned at
the center, with two or three parts of its root system in different
root chambers, or (b) three plants, one in each root chamber.
Each root chamber has its own lines for air inflow (aeration
and stirring), solution inflow, and solution outflow. To reduce
air flows to manageable levels, air fed from a regulated-
pressure supply passes at each chamber through a membrane
placed at the same height, within 5 mm, to prevent siphoning
of solution from one chamber to another after pumping
ceases. Plants are held by electronics-grade, unfilled (translu-
cent), silicone putty around the stem, providing a strong,
nondamaging seal that contains no detectable, leachable traces
of nitrogen- or phosphorous-containing compounds. Each
plant may be removed independently and quickly by remov-
ing the machine screws holding the lid sections together.
The entire three-chamber beaker and its lid are specially
painted to prevent light entry and algal growth and to keep
the temperature within 1°C of air temperature even in full
sunlight and still air. (In contrast, chambers with clean metal
.1
I
Hatched area: One of 3 i
separate root chambers
Figure 2. Top view of lid covering 2-L beaker divided into three
separate root chambers. Lid holds either three plants with undivided
root systems, at locations P1, P2, P3, or one plant with root system
divided into two or three segments, at location PC. Dashed lines
indicate structural features underneath the lid. Details are shown for
only one of the root chambers, for plant at P3.
1128 GUTSCHICK AND KAY
or ordinary white-painted surfaces readily reach 5 to 10°C
above ambient in these conditions). An epoxy-based paint
(Koppers, Inc. item A1737, mil. spec. Mil C 81773, color
17875 white; retail sales through Burbank Paint, Burbank,
CA, and others) reflects shortwave radiation almost com-
pletely, while its very high thermal-infrared emissivity allows
it to lose most of the gained energy. The polystyrene surface
of the breaker must be primed first by painting with a solvent/
roughener (10% by volume powdered chlorinated polyethyl-
ene [Dow Chemical CPE 4213 or 2552] mixed with 80%
tetrahydrofuran and 10% 1,1,l-trichloroethane) and then
with black primer (fast black polyamide epoxy A809C-66,
color 17038 black).
When measuring plant uptake rates or confirming solution
concentrations, we took 1 -L outflow samples from each plant
or treatment inflow, in high-density polyethylene bottles. We
acidified each sample of pH 2 and refrigerated it until analysis.
The concentration depletion times flow rate yielded our esti-
mate of whole-plant uptake, converted as necessary to uptake
rate per unit mass. We analyzed nitrate spectrophotometri-
cally with an Autoanalyzer (cadmium reduction to nitrite and
chromogenic reaction with aromatic reagents [1]). We cor-
rected all peak heights, A, for second-order reaction kinetics
and subsequent nonlinear relation to concentration, c, as A =
bc/(l + ac2); a and b are empirical coefficients depending
upon run conditions (temperature, flow rate, cadmium re-
ductor-column diameter). The parameters a and b are deter-
mined using assays on dilution-series standards. We measured
phosphate spectrophotometrically by the phosphomolybdate-
blue reaction, using ascorbic acid as reductant. For phosphate
concentrations of 1 to 20 Mm (or microformal, to report
properly the summation over ionization states; ref. 11, p. 70),
we extracted the reaction mixture with methyl isobutyl ketone
and read absorbance at 722 nm; this minimized interference
from iron chelates and from temperature shifts. For concen-
trations below 1 MM, we (8) modified a method (6) of triple
extraction of the phosphomolybdate from the aqueous phase
with methyl isobutyl ketone. Only this method proved free
from interference by chelated iron in nutrient solutions.
RESULTS AND DISCUSSION
Our system with seven pumps reliably delivered a variety
of controlled nutrient levels. Over several seasons we have
raised four replicate plants per pump to the stage of 20 g fresh
mass (9) (LE Kay, VP Gutschick, manuscript in preparation;
we began with as many as 12 replicate plants per pump,
sacrificing groups sequentially for physiological analyses.) In
the treatment groups growing at very low nutrient concentra-
tions (0.1 Mm phosphate, 3 Mm nitrate), yet at significant
relative growth rates of 0.08 to 0.10 d-', the nutrient flow rate
required was very high, 400 L d-', and the pumps operated
continuously. We observed neither mortality nor signs of
toxicity from materials (cfJ 10) or from xenobiota. Nutrient
concentrations at the inflow of root chambers held constant
within 2% because the solution mixing ratios were stable.
Concentrations in the root chamber also held constant in the
short term (before plant growth requires adjustment of pump-
ing rates). The 650-mL volume buffers the change in concen-
tration between the existing root-chamber contents and the
Plant Physiol. Vol. 95, 1991
new 95-mL slug of solution delivered on each pump stroke.
One can readily calculate that, if the solution is depleted 30%
from inflow concentration on the average, then an inflow of
1/7 of the chamber volume causes an instantaneous jump of
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6% in nutrient concentration, or only 3% above the mean.
Even at system startup, we observed no transient depression
or elevation of nutrient concentrations, such as adsorption or
desorption from surfaces might cause. For reference, note that
a monolayer of ions on binding sites 0.5 nm apart on the
surface of one root chamber would be equivalent to about 0.3
Mm in the chamber contents of 0.65 L. We attribute this
success partly to materials choice, principally polyethylene,
polystyrene, and surgical-grade Tygon tubing, with limited
exposure of solutions to glass, stainless steel, silicone and
Neoprene rubbers. Furthermore, we cleaned all components
thoroughly with phosphate-free detergents, following this with
acid rinses and final DIW rinses. The solution pH was stable
within 0.3 unit at most, despite variable degrees of nitrate
uptake. Nonrecirculation of solutions aids such pH stability,
while vigorous aeration with air of normal CO2 content
provides good buffering by the carbonic acid-bicarbonate
couple. The flow rates to individual chambers fed by a single
pump were stable individually, while varying as much as 30%
between chambers.
Consequently, we could accurately resolve the time courses
of physiological adaptations in the same plant (LE Kay, VP
Gutschick, manuscript in preparation). These results are re-
ported in preliminary form (9) and will be presented in full
in our companion study, in preparation. Figure 3 shows
2.5
.1-
2.0
._-
a)
1.5
-
1.0
4Q-
* X
0400
4--
0.5
0.0 +
10.0 15.0 20.0 25.0 30.0
Days After Germination
Figure 3. Relationship of root:shoot ratio on a fresh-weight basis to
plant age in Helianthus annuus L. (cv Giant Grey Stripe) as a function
of nominal nutrient levels maintained continuously after day 10 (Mm
nitrate [N] and phosphate [P]). Experiment began with 12 replicate
plants per nutrient treatment group. All plants were germinated in
saturated CaSO4 solution. On d 10 they had attained an initial
root:shoot ratio of 0.39 ± 0.07 on a fresh-weight basis. They were
randomly assigned to one of seven different nutrient treatments
provided by the hydroponic system described herein and grown in a
naturally lit greenhouse with mean day/night temperatures approxi-
mating 320C/200C. Four plants per treatment were harvested on d
17, 22, and 27.
 HYDROPONIC SYSTEM 1129

root:shoot ratio as an example, with (N, P) treatments clearly uct (6). At least 12 replicate plants within any treatment group
differentiated (reanalysis of data of ref. 9, in terms of dry can be monitored for uptake while total equipment cost is
mass). We also resolved modest diurnal cycling in uptake kept modest, by virtue of the flow-division system. The cost
rates (Fig. 4), more so at high nutrient concentrations. We of the entire system is modest compared to that of systems

interpolated these uptake rates and verified that uptake inte-
grated over the whole life of the plant agreed within 10% with
nutrient concentrations in plants.
In addition to the generic advantages (and disadvantages)
of hydroponics, especially allowing repeated, nondestructive
access to functioning roots in situ, our system has several
specific advantages. The nonrecirculation of solutions limits
problems of microbial activity, pH shifts, and cumulative
imbalances in unmonitored nutrients. The system enables
studies of plant performance at very low concentrations,
maintained in steady state or varied according to a chosen
program. We have used 0.07 ,M phosphate and 2.5 lsM nitrate,
at which levels there are no automated on-line methods to
control concentration and calculate uptake, other than costly
flow-injection analysis that can handle relatively few replicate
plants. For example, below several gM, most or perhaps all
high-resolution phosphate assays are batch methods using
multiple solvent extraction of phosphomolybdate assay prod-
1.25
/ .p \ ...
_ / /~ N\..o :f
1:3
-
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using peristaltic pumps and automatic monitoring and
makeup of nutrients. The cost is low particularly if the re-
searcher can machine the components, using a milling ma-
chine, drill press, and welder. The reciprocating pumps are
cost-effective in recirculating systems also. One can also main-
tain disjoint nutrient concentrations on two or three separate
root system segments of individual plants while monitoring
their uptake rates via grab-samples of inflow and outflow
solutions. Finally, we note the excellent control of heat gain
from shortwave radiation absorption, afforded by the special
white epoxy paint. This paint is also useful in shielding air
temperature sensors and the like.
Several disadvantages accrue from nonrecirculation of the
solutions. The high volume of DIW and nutrient use incurs
a significant operational expense. We used 15,000 L in one
2-week experiment on 84 plants, 28 of which grew to 14 g
average fresh mass. (Recirculating systems also require dis-
carding of large reservoirs after an experiment, as much as
2000 L [3]). Such expense also precludes the use of PEG for
osmotic stress studies. The large volumes for disposal may
preclude most uses of radioactive tracers, although we used
33P at very low levels within radiation-safety guidelines. The
researcher must monitor pump rates and pump function
frequently. One must also run off-line solution analyses every
several days to assure that pumping rates are sufficient.

0
1.00
/
// /~
/// N\ : '' '''''
' ACKNOWLEDGMENTS
We wish to thank a number of people at Los Alamos National
/ // ' '' \ Laboratory for their help. Faustin Trujillo and Dan Talley helped in
-4- the design and initial machining of the pump components. Michael
co
// // ~ \'- A. Wolf and David Waechter helped in the design and construction,
;., 0.75 respectively, of the electronic controls. Consuelo Montoya helped us
operate the system at times. James Steger, Caroline Reynolds, and
N-N 25- Ralph Franklin (U.S. Department of Energy; currently at Clemson
University) provided critical administrative support.
_ / /N10P.N
/ N LITERATURE CITED
> 0.50
cu
/ -N40P. 1. Armstrong FAJ, Sterns CR, Strickland JDH (1967) The meas-
-N.....P.8 urement of upwelling and subsequent biological processes by
CZ 0.25
co
means of the Technicon AutoAnalyzer and associated equip-
ment. Deep-Sea Res 14: 381-389
025 2. Asher CJ, Edwards DG (1983) Modern solution culture tech-
niques. In A Lauchli, RL Bieleski, eds, Encyclopedia of Plant
Pb~~~ .0 .O.R.Ov. Physiology (New Series, Vol 15A). Springer-Verlag, Berlin, pp
95-1 18
3. Asher CJ, Ozanne PG, Loneragan JF (1965) A method for
controlling the ionic environment of plant roots. Soil Sci 100:
149-156
4. Bloom AJ (1989) Continuous and steady-state nutrient absorp-
tion by intact plants. In JG Torrey, LW Winship, eds, Appli-
10.0 12.5 15.0 17.5 20.0 22.5 25.0 cations of Continuous and Steady-State Methods to Root
Time After Midnight, 11 October [h] Biology. Kluwer Academic, Dordrecht, pp 147-163
5. Breeze VG, Canaway RJ, Wild A, Hopper MJ, Jones LHP
Figure 4. Instantaneous rate of phosphate uptake per unit fresh (1982) The uptake of phosphate by plants from flowing nu-
mass of plant, as a function of time in diurnal cycle, for plants at end trient solution. I. Control of phosphate concentration in solu-
tion. J Exp Bot 33: 183-189
of experiment (8) that generated the data in Figure 1. Uptake rates 6. Gibson AR, Bailey JM, Giltrap DJ (1976) Determination of
are resolved by nutrient treatment during growth and for individual trace amounts of phosphate in water extracts of soil. Commun
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phosphate in demineralized and demineralized-distilled water.
Talanta 32: 93-94 12.
9. Gutschick VP, Kay LE (1986) Root adaptations at stress levels 13.
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