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HORTSCIENCE 39(6):12871292. 2004.

environmental variable affecting WUE, and


therefore possibly optimal fertilizer con-
Light Intensity and Fertilizer centration of subirrigated plants. Water-use
efficiency normally increases with increasing
light intensity (Alexander and Donnelly, 1995;
Concentration: I. Estimating Optimal Caviglia and Sadras, 2001; Israeli et al., 1996;
Le Roux et al., 2001; Ponton et al., 2002;
Fertilizer Concentrations from Vandana, 1999). Photosynthetic photon flux
varies considerably across the country and
Water-use Efficiency of Wax Begonia throughout the year. Thus, fertilizer recom-
mendations may have to be adjusted based
Krishna S. Nemali and Marc W. van Iersel1 on the prevailing PPF level. Surprisingly,
research on optimum fertilizer concentrations
Department of Horticulture, Miller Plant Sciences Building, The University of plants grown under different PPF levels
of Georgia, Athens, GA 30602-7273 is scant, although there is some research that
Additional index words. Begonia semperflorens-cultorum , electrical conductivity, nutrient looked at the effect of other environmental
factors on nutrient uptake. Kang and van Iersel
uptake, photosynthetic photon flux, tissue nutrient analysis
(2001) concluded that temperature affects the
Abstract. Environmental conditions and incorporation of nutrients into the growing medium optimal fertilizer concentration of subirrigated
can affect the fertilizer needs of bedding plants. To evaluate the effects of photosynthetic petunias (Petunia hybrida Hort. Vilm-Andr.),
photon flux (PPF) and starter fertilizer on the fertilizer requirements of subirrigated plants, whereas Gislerrd and Mortensen (1990) found
we grew wax begonias (Begonia semperflorens-cultorum Hort.) under three PPF levels (aver- that tissue nutrient concentration of Begonia
aging 4.4, 6.2, and 9.9 molm2d1) and four fertilizer concentrations [electrical conductivity heimalis was lower at high (90%) than a
(EC) of 0.15, 0.33, 0.86, and 1.4 dSm1] in a normal (with starter fertilizer, EC = 2.1 dSm1) low (60%) relative humidity, but increased
and heavily leached (with little starter fertilizer, EC = 0.9 dSm1) growing medium. Except when the concentration of nutrient solution
for shoot dry mass, we did not find any significant interactions between PPF and fertilizer was increased.
concentration on any of the growth parameters. There was an interactive effect of fertilizer The objective of this experiment was to
concentration and starter fertilizer on all growth parameters (shoot dry mass, leaf area, plant study the effect of starter fertilizer and varying
height, and number of flowers). When the growing medium contained a starter fertilizer, light intensities on optimal N concentration
fertilizer concentration had little effect on growth. When the growing medium was leached of the fertilizer solution for the production of
before transplanting, growth was best with a fertilizer EC of 0.86 or 1.4 dSm1. Water-use subirrigated begonias. We hypothesized that
efficiency (WUE) was calculated from 24-hour carbon exchange and evapotranspiration the optimal N concentration would increase
measurements, and used to estimate the required [N] in the fertilizer solution to achieve a with increasing PPF (because of the increase
target tissue N concentration of 45 mgg1. Increasing PPF increased WUE and the required in WUE) and decrease with a starter fertil-
[N] (from 157 to 203 mgL1 at PPF levels of 4.4 and 9.9 molm2d1, respectively). The PPF izer in the growing medium. Wax begonia
effect on the required [N] appeared to be too small to be of practical significance, since dry was used as a model crop, because it can be
mass data did not confirm that plants grown at high light needed higher fertilizer concentra- grown under a variety of light intensities, and
tions. Thus, fertilizer concentrations need not be adjusted based on PPF. it is one of the most popular shade-tolerant
plants in floriculture.
Subirrigation systems have become more the amount of nutrients supplied to the plants
popular in recent years due to zero runoff, and (nutrient supply). To prevent nutrient deficien- Materials and Methods
increased efficiency in fertilizer and water cies or toxicities, nutrient supply and demand
use (Elliot, 1990; Morvant et al., 1997; Uva should be balanced. The volume of the applied Plant material. Plug seedlings of wax
et al., 1998; van Iersel, 1996; Yelanich and fertilizer solution absorbed by the growing me- begonia Cocktail Vodka were obtained
Biernbaum,1990). Because of the absence of dium equals evapotranspiration, if there is no from a commercial grower (Speedling Inc.,
leaching in subirrigation, starter fertilizer can leaching and the medium is rewatered to the Blairsville, Ga.) and transplanted into square,
remain in the growing medium for a longer same moisture level at each irrigation. Thus, 10-cm (510-mL) containers filled with a soil-
period, and reduce the fertilizer requirements to maintain the balance between supply and less growing medium (Fafard 2P mix; Fafard,
by supplying a substantial amount of nutrients demand for N nutrition specifically Anderson, S.C.) on 13 June 2001. The starter
to plants. Water-use efficiency (WUE, the [Ntissue] GR = [Nfert] ET [Eq. 1] fertilizer in the growing medium contained 89,
ratio of growth to transpiration in plants) can where [Ntissue] is the desired tissue N concen- 37, and 32 gm3 of N, P, and K, respectively.
also affect the fertilizer requirements of plants tration (mgg1), GR is the growth rate (gd1), The starter fertilizer was leached out of half
(Bugbee, 1995). A plant with a high growth to [Nfert] is the required N concentration of the of the pots by watering heavily for about ten
transpiration ratio takes up a relatively small fertilizer solution to maintain the balance be- times over a three day period. The EC of the
amount of water while producing a gram of dry tween N supply and demand (mgL1), and ET normal (hereafter, with starter fertilizer) and
matter. Therefore, a higher fertilizer concentra- is evapotranspiration (Ld1). Since GR/ET is leached (hereafter, without starter fertilizer)
tion should be supplied to these plants to supply the WUE: growing medium were determined with the
adequate nutrients. [Nfert] = [Ntissue] WUE [Eq. 2]. pour-through technique (Wright, 1986) to
The product of tissue nutrient concentration This concept has been used in hydroponic be 2.1 and 0.9 dSm1, respectively. After
and the mass of newly formed tissue determines studies, where photosynthesis and transpiration transplanting, the seedlings were placed on
the amount of nutrients taken up by the plant models can be used to predict the optimal electri- 1.2 2.4-m2 ebb-and-flow benches (Midwest
(nutrient demand), while the product of the cal conductivity (EC) of the supplied nutrient GroMaster, St. Charles, Ill.) and subirrigated
volume of the applied fertilizer solution and solution (Klring and Cierpinski, 1998). In daily with 20N4.4P16.6K fertilizer solu-
the nutrient concentration of the solution equals bedding plant production, evaporation from tions (Peters 201020 peat-lite special,
the growing medium may be an important fac- The Scotts Co., Marysville, Ohio). Fertilizer
Received for publication 30 Jan. 2003. Accepted for tor in determining the total water-use, and the solutions were stored in plastic barrels (210
publication 3 Sept. 2003. We thank Keven Calhoun combined value of evaporation from the grow- L) and pumped into the watertight trays of
and Larry Freeman for their technical assistance and ing medium and transpiration from the plants the ebb-and-flow system using submersible
Svoboda Pennisi and Hugh Earl for their helpful should be used to calculate optimal fertilizer pumps (NoKorode-2; Little Giant, Oklahoma
comments on earlier drafts of this manuscript. City, Okla.). The bottoms of the pots were im-
1
Corresponding author; e-mail mvanieruga.edu.
concentrations.
Photosynthetic photon flux is an important mersed in fertilizer solution for about 13 min

HORTSCIENCE VOL. 39(6) OCTOBER 2004 1287


(5 min for pumping and 8 min for draining) growing medium, which in turn may affect the complete block with a split-split-plot design
during which the growing medium absorbed subsequent growth of the plants. and three replications. Fertilizer EC was the
it by capillary action. The EC of the fertilizer Only plants grown without a starter fertilizer main blocking factor, with three light intensi-
solution was measured using an EC meter in the growing medium were used in the WUE ties as main splits, and a group of 30 plants
(model M90, Corning, Corning, N.Y.) and study. Four plants from each experimental unit either with or without starter fertilizer was a
adjusted weekly by adding fertilizer when the were placed in an eight-chamber, whole-plant subsplit (experimental unit). Data collected
barrels were refilled. Plants were grown in a gas exchange system (van Iersel and Bugbee, on physical growth parameters, including dry
greenhouse covered with double-layered poly- 2000) at the end of the experiment to measure mass, leaf area, plant height, and number of
ethylene and a shade cloth which transmitted the CO2 exchange rates. Since there were flowers per plants, and EC and pH of the grow-
63% of the incident light. The average day 36 experimental units [3 replications 12 ing medium were subjected to ANOVA and
and nighttime temperatures were 32.2 and 23.7 treatments (4EC 3 PPF levels)], and eight regression analysis using statistical analysis
C, respectively (with an average of 27.1 C groups of plants could be measured simultane- software (SAS institute, Cary, N.C.). Since
over the course of the experiment). ously, these data were collected during a 5-d the absence or presence of starter fertilizer is a
Treatments. Plants were placed on one of period. Plants were kept in the gas exchange noncontinuous variable (i.e., a class variable),
12 ebb-and-flow benches, and each bench chambers for a period of 24 h (14 h of light, it could not be included in regression models.
was subirrigated with one of four fertilizer 10 h of dark), during which whole-plant net Plant growth was modeled as a function of
concentrations (0, 30, 110, or 190 mgL1 N, photosynthesis and dark respiration (Pn and EC according to:
corresponding ECs of 0.15, 0.33 , 0.86, and 1.4 Rd respectively, expressed in mols1) rates Y = 0 + 1 ln(EC) 2 EC [Eq. 5]
dSm1, respectively). Each bench was divided were measured once every 10 min. Inside the where 0 ...2 are regression coefficients. The
into three sections with different PPF levels, growth chambers, plants were exposed to the log transformation of EC was included, because
by placing different shade cloths over PVC same PPF as in the greenhouse. To determine it resulted in a good empirical fit, while simple
frames, which were placed on each bench. This ET, pots were weighed before and after the gas linear or quadratic regression did not describe
resulted in 63%, 35%, or 0% shade (in addition exchange measurements. Growth rate (GR, the data adequately. Since there was a signifi-
to the shade due to the shade cloth covering gd1; amount of dry matter produced by a cant two-way interactive effect of fertilizer EC
the greenhouse), equivalent to an average daily group of four plants in a day) was calculated and PPF on shoot dry mass, regression were
PPF of 4.4, 6.2, and 9.9 molm2d1, respec- as follows: done separately for each PPF level. Since the
tively. Two groups of 30 plants each (with or GR = [(Pn,avg tlight Rd,avg tdark) 12/fC ] [Eq. 3] interaction between PPF and the presence of
without starter fertilizer), were grown in each where Pn,avg and Rd,avg are the average net pho- starter fertilizer was not significant for any of
bench section with different PPF levels. Six tosynthesis and dark respiration rates (mols1), the growth parameters, data were averaged over
quantum sensors (QSO-SUN; Apogee Instru- tight and tdark are the durations of light and dark all three PPF levels to describe the effect of
ments, Logan, Utah) were arranged in different periods in seconds, 12 converts moles of carbon fertilizer EC on plant growth in the presence
experimental units to measure daily PPF. to grams of carbon, and fC is the carbon con- and absence of starter fertilizer. A log transfor-
By watering daily, plants were provided tent (1/fc converts grams of carbon to grams of mation was not required in the analysis of the
with enough water to prevent drought stress in dry matter) determined from total shoot tissue EC of the growing medium, since the trends
any of the treatments. Although all plants were analysis of plants in different treatments (fc could be adequately described by simple linear
subirrigated daily, this does not imply that all ranged from 0.39 to 0.43). Water-use efficiency regression.
plants received equal amounts of water. With (gL1) was calculated as The analysis of physiological parameters
subirrigation, the growing medium absorbs wa- WUE = GR/ET [Eq. 4] (net photosynthesis, dark respiration, evapo-
ter to near saturation, and the actual amount Optimal nitrogen concentrations of the fer- transpiration, growth rate, WUE, and [Nfert])
of water, and thus fertilizer, absorbed by the tilizer solutions were calculated as described in was done using multiple regression, with an
growing medium depends on the ET since the Eq. 2, assuming a desired [Ntissue] of 45 mgg1, interaction term:
last irrigation. Thus, the growing medium of based on the recommended range of 20 to 60 Y = Y0 + Y1 EC + Y2 PPF + Y3 EC2 +
plants with high ET rates (i.e., larger plants mgg1 (Mills and Jones, 1996). Note that the Y4 PPF2+ Y5 EC PPF [Eq. 6]
grown under higher PPF) absorbed more water assumed value for [Ntissue] will affect the cal- where Y0.....Y5 are regression coefficients.
and fertilizer than plants with low ET rates. culated [Nfert], but not the relative differences Nonsignificant terms were removed by back-
Measurements. Data on plant height, leaf among treatments. Those differences are solely ward selection (P < 0.05).
area, shoot dry mass, and number of flow- dependent on WUE instead. A similar regression was done with [Nfert] as
ers were collected at the end of the growing The entire shoot samples were analyzed the dependent variable, except that the actual
period (6 weeks after transplanting). Plant to determine the tissue nutrient concentration. N concentration of the fertilizer solution ([N]a)
height was measured from the surface of Tissue C, N, and S concentrations were mea- was used as the independent variable, instead
the growing medium to the top of the plant. sured using a CNS analyzer (model 2000; of EC. If [N]a is lower than [Nfert], either the
Leaf area was measured using an area meter Leco corporation, St. Joseph, Mich.) and the actual N concentration in the plant will be
(LI-3100; LI-COR, Lincoln, Nebr.). Shoots other nutrients were measured using a ICAP lower than [Ntissue], or the [N] in the growing
from sample plants were dried in a forced-air analyzer (model 9000; Thermo Jarrell Ash medium will decrease. Vice versa, if [N]a is
oven maintained at 80 C for 1 week before corporation, Franklin, Mass.). higher than [Nfert], this will result in a tissue
measuring their dry mass. Flowers from six From the collected data, [Nfert] was calcu- [N] higher than 45 mgg1, or an increase in
plants in each experimental unit were counted. lated as follows: if Pn,avg (during the 14-h light the [N] of the growing medium. The [N] in
Electrical conductivity and pH of growing period) and Rd,avg (during the 10-h dark period) the fertilizer solution to maintain the balance
medium were collected three times during rates of plants were 1.0 and 0.5 mols1, respec- between supply and demand (assuming 45
the growing period (after 2, 4, and 6 weeks), tively, and fc of the plants was 0.40, then GR of mg.g1 N in the tissue) can be determined as
using the pour-through technique (Wright, plants can be calculated as GR = {[(1.0 14 the concentration at which [N]a and [N]fert are
1986). About 25 mL of tap water (EC of 0.1 3600) (0.5 10 3600)]/1,000,000} 12/ equal (or [N]a [N]fert = 0).
to 0.2 dS.m1) was poured evenly on top of the 0.4 = 0.972 gd1. The denominator 1,000,000
growing medium an hour after subirrigating converts the photosynthetic or respiration rate Results and Discussion
the plants, and the collected leachate was used from mols1 to mols1. Let us assume that
to measure EC and pH of the growing medium 200 mL of water is lost in evapotranspiration in Plant growthlight intensity relationship.
with EC and pH sensors (model M90; Corning, producing 0.972 g of dry matter during the 24- The three-way interaction among PPF, fertil-
Corning, N.Y.). Plants used for determining EC h period. Therefore, WUE = 0.972/0.2 = 4.86 izer EC, and starter fertilizer on shoot dry mass
of the growing medium were not used in any gL1 , and [Nfert] = 4.86 45 = 219 mgL1. was nonsignificant (P = 0.07), but there was
subsequent analyses, because the pour-through Experimental design and analysis. The an interactive effect of PPF and fertilizer EC
method may change salt gradients within the treatments were organized in a randomized on shoot dry mass, indicating that the growth

1288 HORTSCIENCE VOL. 39(6) OCTOBER 2004


dry mass (360%), leaf area < 0, not enough N is applied to maintain tissue
(425%), plant height (480%), [N] at 45 mg.g1 unless the [N] in the growing
and flower number (280%) medium decreases, and when [N]a [Nfert] >
when the irrigation solution 0, excess N will accumulate either in the plant
did not contain fertilizer (EC or the growing medium. Supply and demand
= 0.15 dSm1), but had little for N are balanced when [N]a [Nfert] = 0.
effect on plant growth if the Photosynthetic photon flux affected at which
irrigation solution did con- [N]a this occurred. At PPF levels of 4.4, 6.2,
tain fertilizer (Fig. 2). Since and 9.9 molm2d1, supply and demand were
even the highest fertilizer EC equal at [N]a of 157, 172, and 203 mgL1 N,
did not result in a clear reduc- respectively.
tion in plant growth, optimal This suggests that fertilizer concentrations
fertilizer concentrations should be increased with increasing PPF. How-
cannot reliably be estimated ever, this conclusion was not confirmed by the
from these data. Nemali and dry mass data, since even the highest fertilizer
van Iersel (2004) found that EC did not result in a decrease in dry mass.
a fertilizer EC of 1.3 dSm1 This made it impossible to estimate the optimal
resulted in maximum growth fertilizer EC from the dry mass data (Fig. 1).
Fig. 1. Effect of fertilizer electrical conductivity of wax begonias in a growing medium without There was no interactive effect of fertilizer EC
(ECfert) and daily photosynthetic photon flux on any starter fertilizer, irrespective of PPF. and PPF on leaf area, indicating that fertilizer
shoot dry mass of subirrigated wax begonias at Light intensity WUE and [Nfert] relation- EC effects on leaf area were similar at different
6 weeks after transplanting. The error bar indi- PPF levels. In a subsequent, similar experiment
ships. The fitted polynomial equation (Eq. 6)
cates the interactive least significant difference
among fertilizer ECs within one light intensity indicated that there was an interactive effect with higher fertilizer concentrations, including
and among light intensities within one fertilizer of fertilizer EC (or [N]) and light intensity on superoptimal ones, we did not find an effect of
EC. The lines indicate significant effects (P < WUE and [Nfert] of plants. These models indi- PPF on the [N] of the fertilizer solution result-
0.05); DWshoot = 1.62 + 0.30 ln(ECfert)) (r2 = cate that both WUE and [Nfert] increased with ing in maximal growth (Nemali and van Iersel,
0.70) and 3.47 + 1.12 ln(ECfert) + 1.23 EC increasing PPF, especially at higher fertilizer 2004). Apparently, the effect of PPF on the
(R2 = 0.83), for medium (6.2 molm2d1) and concentrations (Fig. 3, Table 1). Earlier studies optimal [N] of the fertilizer is too small to have
high PPF (9.9 molm2d1), respectively. also reported an increase in WUE of various a practical impact. One possible reason for the
species with increasing PPF (Alexander et al., lack of a practical effect of PPF on the optimal
response to fertilizer EC depended on the PPF 1995; Caviglia and Sadras, 2001; Israeli et al., [N] is that plants can perform well over a wide
level. Irrespective of fertilizer EC, shoot dry 1996; Le Roux et al., 2001; Ponton et al., 2002; range of tissue [N]. Increased leaf [N] does not
mass increased with increasing PPF (Fig. 1). Vandana, 1999). Increasing fertilizer EC from always contribute to photosynthetic N, but may
At all three PPF levels, dry mass increased as 0.15 to 0.86 dSm1 also increased WUE and contribute to stored N, which can be used at
the fertilizer EC was increased from 0.15 to thus [Nfert], while a further increase in fertilizer times of a N deficiency (Thornley, 1995). It
0.33 dSm1. This increase in dry mass with EC resulted in a small decrease (Fig. 3). is possible that tissue [N] < 45 mgg1 would
increasing EC was more pronounced at medium There generally was a difference between have been sufficient as well, since the optimal
or high (6.2 or 9.6 molm2d1) than at low (4.4 [N]a and [Nfert] (Fig. 4), indicating that N supply range for begonia reportedly is 20 to 60 mgg1
molm2d1) PPF, presumably because PPF, and demand were not balanced in such a way (Mills and Jones, 1996). In addition, the grow-
not nutrition, was the main limiting factor for to maintain N in the plants. When [N]a [Nfert] ing medium can act as a buffer for nutrients,
growth at low PPF. Similarly, Larouche et
al. (1989) concluded that vegetative growth
of greenhouse tomatoes (Lycopersicon escu-
lentum Mill. Vedettos) was limited by light
at low PPF, and therefore did not respond to
increased N in the nutrient solution. However,
we did not find any significant interaction
between PPF and fertilizer EC on leaf area,
plant height, or number of flowers per plant.
Irrespective of fertilizer EC and starter fertil-
izer, these variables increased linearly with
increasing PPF (data not shown).
Plant growthstarter fertilizer relationship.
There were interactive effects of fertilizer EC
and starter fertilizer on shoot dry mass, leaf
area, plant height, and number of flowers per
plant (Fig. 2). These interactions indicate
that, irrespective of PPF level, plant growth
response to fertilizer EC depended on the pres-
ence or absence of starter fertilizer. When the
growing medium contained starter fertilizer,
shoot dry mass and number of flowers did not
respond to increasing fertilizer EC, while leaf
area and plant height increased slightly with Fig. 2. Effect of fertilizer electrical conductivity (EC) and starter fertilizer level on shoot dry mass, leaf area
an increase in fertilizer EC from 0.15 to 0.33 (LA), plant height (PlHt), and flower number of subirrigated wax begonias at 6 weeks after transplanting.
dSm1 and decreased with a further increase Error bars indicate the interactive least significance difference (LSD0.05) between fertilizer ECs within one
in fertilizer EC. There was a strong increase starter fertilizer level and between starter fertilizer levels within one fertilizer EC. The lines indicate
in shoot dry mass, leaf area, and plant height significant effects (P < 0.05). Without a starter fertilizer, DWshoot = 2.802 + 1.153 ln(EC) 1.153
in response to increasing fertilizer EC in the EC, (R2 = 0.88), LA= 0.15 + 0.064 ln(EC) 0.076 EC, (R2 = 0.83), Pl Ht = 38.69 + 17.12 ln(EC)
absence of a starter fertilizer. Starter fertilizer 20.55 EC, (R2 = 0.97) and with a starter fertilizer, LA = 0.11 + 0.018 ln(EC) 0.036 EC, (R2
in the growing medium greatly increased shoot = 0.58) and PlHt = 28.93 + 6.75 ln(EC) 12.44 EC, (R2 = 0.52).

HORTSCIENCE VOL. 39(6) OCTOBER 2004 1289


in different treatments and growing medium contained a starter fertilizer,
the relationship among ET, this suggests that even a growing medium EC
PPF, and fertilizer EC was of 0.64 dSm1 may be sufficient for begonia.
poor (R2 = 0.37). In general, However, when the growing medium contained
ET increased with increasing a starter fertilizer and the fertilizer EC was 0.15
PPF and also with increasing dSm1 (i.e., no fertilizer in the solution, Fig.
fertilizer EC. 5), the EC of the growing medium decreased
EC and pH of the growing during the experiment. Since the pour-through
medium. There was an interac- method measures the EC in the bottom part of the
tive effect of fertilizer EC and growing medium, this decrease in EC indicates
starter fertilizer on growing that either the plants removed nutrients from
medium EC at 4 and 6 weeks the growing medium during the experiment or
after transplanting, but not at that nutrients moved to top layer of the grow-
2 weeks after transplanting ing medium, as can happen with subirrigation
(Fig. 5). Electrical conductiv- (Nemali and van Iersel, 2004).
ity of the growing medium In the presence of a starter fertilizer in
increased with increasing the growing medium, plants grew best with a
fertilizer EC, and was always fertilizer EC of 0.33 dS.m1, while the optimal
Fig. 3. Effect of photosynthetic photon flux and fertil- higher with than without a starter fertilizer. At 4 fertilizer EC was 0.86 to 1.40 dSm1 in the
izer concentration (EC) on water-use efficiency and 6 weeks after transplanting, the increase in absence of starter fertilizer. This corresponds to
(WUE) of subirrigated wax begonia at 6 weeks growing medium EC with increasing fertilizer a final growing medium EC of 1 to 2.5 dSm1.
after transplanting. WUE = 0.62 + 9.33 EC
EC was higher for growing medium with than However, since there was no clear reduction
6.11 EC2 + 0.32 EC PPF, R2 = 0.78.
without a starter fertilizer. Electrical conductiv- in growth at the highest EC, a reliable upper
ity of the growing medium was not affected by EC limit for adequate growth could not be
and changes in the nutrient concentrations in PPF. This supports our finding that the fertilizer established. However, in subsequent research
the growing medium do not necessarily affect concentration for subirrigated wax begonia need we found that a growing medium EC of 1.4 to
tissue nutrient concentrations. For example, if not be changed with changing light intensities. 2.8 dSm1 results in optimal growth (Nemali
N supply is less than demand, it is possible that It is important to realize that the lack of PPF and van Iersel, 2004). In contrast, James and
tissue [N] is relatively stable while [N] in the effects on growing medium EC does not imply van Iersel (2001), reported that plant growth
growing medium decreases. that PPF did not affect nutrient uptake. Plants of wax begonias was not reduced by more than
Gas exchange parameters. There was an grown at higher PPF had more evapotranspira- 10 percent when the growing medium EC was
interactive effect of PPF and fertilizer EC on Pn tion (Table 1), and the growing medium of these increased from 2.1 to 6.3 dS.m1.
and Rd of the plants (Table 1). There was little plants thus absorbed more water and fertilizer Growing medium pH was not affected by
or no effect of PPF on Pn and Rd in unfertilized during each subirrigation event. light intensity, but decreased with increasing
treatments (EC = 0.15 dSm1), but both Pn and The EC of growing medium at the end of fertilizer EC due to the acid-forming nature of
Rd increased linearly with PPF in fertilized the experiment (week 6, Fig. 5) ranged from the fertilizer. When the growing medium con-
treatments, and this increase was greater at 0.64 to 3.65 dSm1 with starter fertilizer and tained a starter fertilizer, the pH values ranged
higher fertilizer EC (i.e. a positive fertilizer 0.32 to 2.4 dS.m1 without a starter fertilizer. from 5.2 to 6.2 and without a starter fertilizer
EC PPF interaction term). Both Pn and Rd James and van Iersel (2001) found that growth from 5.3 to 6.8 (data not shown). These values
increased as fertilizer EC increased from 0.15 of wax begonias was acceptable when the EC are in or near the recommended range (5.5 to
to 0.86 dSm1, with little or no further increase of the growing medium remained within a 6.5) for most greenhouse crops (Lang, 1996).
as fertilizer EC increased from 0.86 to 1.40 range of 2.1 to 5.4 dSm1. Thus, wax begonias When irrigated with tap water (0.15 dS.m1), pH
dSm1. Similarly, van Iersel and Kang (2002) can tolerate higher growing medium ECs than of the growing medium did not differ signifi-
reported a quadratic relationship between Pn and occurred in this experiment, which explains cantly among plants grown with and without a
Rd and increasing fertilizer EC in pansy (Viola the lack of a decrease in growth at the highest starter fertilizer. This suggests that lime was not
wittrockiana Gams.). There was no interactive fertilizer EC (Figs. 1 and 2). Since we did not see completely lost in leaching the starter fertilizer
effect of PPF and fertilizer EC on ET of plants large effects of fertilizer EC on growth when the from the growing medium.
Table 1. Effect of fertilizer electrical conductivity (EC) and photosynthetic photon flux (PPF) on dark respiration (Rd), net photosynthesis (Pn), evapotranspiration
(ET), and growth rate (GR) of subirrigated wax begonia. Data were collected at 6 weeks after transplanting. Regression coefficients are based on Y = Y0 +
Y1 EC + Y2 PPF + Y3 EC2 + Y4 PPF2 + Y5 EC PPF.
Fertilizer EC PPF Rd Pn ET GR
(dSm1) (molm2.d1) (mols1) (mols1) (mLd1) (gd1)
0.15 4.4 0.03 0.05 63 0.04
6.2 0.04 0.08 71 0.07
9.9 0.05 0.06 123 0.04
0.33 4.4 0.09 0.31 146 0.35
6.2 0.09 0.44 162 0.55
9.9 0.13 0.52 190 0.62
0.86 4.4 0.09 0.41 139 0.53
6.2 0.12 0.54 168 0.71
9.9 0.21 0.76 179 0.92
1.40 4.4 0.10 0.30 162 0.34
6.2 0.16 0.55 193 0.66
9.9 0.21 0.77 200 0.94
Regression coefficients
R2 0.67 0.81 0.37 0.81
Intercept 0.05*** 0.07NS 54.7 0.13
EC NS
0.98*** 58.16* 1.37***
EC2 0.032* 0.707*** --- 0.97***
PPF --- --- 8.01* ---
EC PPF 0.018*** 0.063*** --- 0.08***
NS,*,**,***
Nonsignificant or significant at P < 0.05, 0.005, or 0.0005, respectively.

1290 HORTSCIENCE VOL. 39(6) OCTOBER 2004


Fig. 4. The effect of photosynthetic photon flux (PPF) and fertilizer concentra-
tion of the fertilizer ([N]a) on the difference between [N]a and the fertilizer
concentration needed to maintain tissue [N] at 45 mgg1 ([Nfert]). When [N]a
[Nfert] < 0, N supply is less than demand, when [N]a [Nfert] > 0, supply
exceeds demand, and when [N]a [Nfert] = 0 supply equals demand. Arrows
indicate the [N]a at which N supply and demand are balanced at different
PPF levels. Curves were calculated from the regression of [Nfert] versus [N]a:
[Nfert] = 40.65 1.12 [N]a + 0.0119 [N]a2 + 0.1096 [N]a PPF, R2 =
0.75. Data points are the mean of three replications.

Tissue nutrient composition. Tissue N, Fe, Fig. 5. Effect of fertilizer


and Zn concentrations responded quadratically electrical conductivity
to increasing fertilizer EC, both in the presence (ECfert) and starter fer-
and absence of a starter fertilizer. However, tilizer on the EC of the
growing medium (ECmed) at 2, 4, and 6 weeks after transplanting. Error bars
tissue K concentration responded quadrati- represent the interactive least significant difference (LSD0.05) between fertilizer
cally with increasing fertilizer EC only in the concentrations within one starter fertilizer level, and between starter fertilizer
absence of a starter fertilizer, while there was levels within one fertilizer concentration. The lines indicate significant cor-
no effect of fertilizer EC on K in the presence relations (P < 0.05). Week 2: ECmed = 1.29 + 0.86 ECfert , r2 = 0.89 (with
of starter fertilizer (Table 2). There was no starter) and ECmed = 0.38 + 0.86 ECfert, r2 = 0.89 (without starter). Week
response of tissue P, Ca, Mg, S, Cu, Mo, and 4: ECmed = 0.54 + 1.35 ECfert, r2 = 0.90, (with starter fertilizer) and 0.07 +
Al concentrations to increasing fertilizer EC, 1.15 EC fert, r2 = 0.98, (without starter fertilizer). Week 6: EC med = 0.23 +
either in the presence and absence of a starter 2.42 ECfert, r2 = 0.89, (with starter fertilizer) and 0.01 + 1.62 ECfert, r2 =
fertilizer. Overall, PPF had no effect on tissue 0.79 (without starter fertilizer).
nutrient composition, except for tissue Na, B,
and Mn concentrations, which responded qua- at high light intensity should be grown with 1.4 dS.m1, while lower concentrations (EC <
dratically to increasing PPF in the presence of higher fertilizer concentrations. However 0.86 dSm1) would be sufficient for growing
a starter fertilizer. this effect may be too small to be of practical media with a starter fertilizer.
significance, because this finding could not be
Conclusions confirmed based on dry mass or leaf area data. Literature Cited
Based on our findings, wax begonias grown Alexander, J.D. and J.R. Donnelly. 1995. Photosyn-
The effect of increasing PPF and fertilizer without a starter fertilizer should be fertilized thetic and transpirational responses of red spruce
EC on WUE and [Nfert] suggest that plants with a fertilizer solution with an EC of 0.86 to under-storey trees to light and temperature. Tree
Table 2. Regression analysis of effect of starter fertilizer, fertilizer electrical conductivity (EC), and photosynthetic photon flux (PPF) on tissue nutrient concentration
of subirrigated wax begonia at the end of the experiment. Regression coefficients are based on Y = Y0 + Y1 EC + Y2 PPF + Y3 EC2 + Y4 PPF2. There
were no significant interactive effects between EC and PPF on any of the nutrients. In the absence of EC or PPF effects, the intercept represents the average
concentration from all treatments combined.
N P K Mg S Ca Na Cu Mo Al B Fe Mn Zn
Parameter (mgg1) (gg1)
With starter
R2 0.91 --- --- --- --- --- 0.92 --- --- --- 0.91 0.83 0.68 0.69
Intercept 50NS 5.4 44.8 7.7 3.0 8.8 5.1*** 8.3 4.4 108 64** 168* 598** 23NS
EC 59* --- --- --- --- --- --- --- --- --- --- 90* --- 64*
PPF NS --- --- --- --- --- 0.7* --- --- --- 11* --- 112* ---
EC2 32* --- --- --- --- --- --- --- --- --- --- 62* --- 39*
PPF2 --- --- --- --- --- --- 0.05* --- --- --- 0.69* --- 7.2* ---
Without starter
R2 0.89 --- 0.94 --- --- --- --- --- --- --- --- 0.65 --- 0.89
Intercept 36NS 5.0 17NS 8.6 2.9 10.0 2.2 8.5 7.4 116 31 80NS 244 46*
* **
EC 87 --- 58 --- --- --- --- --- --- --- --- --- --- ---
PPF NS --- --- --- --- --- --- --- --- --- --- --- --- ---
EC2 43* --- 33* --- --- --- --- --- --- --- --- 74.1* --- 18*
PPF2 --- --- --- --- --- --- --- --- --- --- --- --- --- ---
NS,*,**,***
Nonsignificant or significant at P < 0.05, 0.005, or 0.0005, respectively.

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Physiol. 15: 393398. Kang, J.G. and M.W. van Iersel. 2001. Interactions differing in light requirement and growth rate.
Bugbee, B. 1995. Nutrient management in recirculat- between temperature and fertilizer concentration HortScience 39(6):12951299.
ing hydroponic culture, p. 1530. In: Proc 16th affect growth of subirrigated petunias. J. Plant Ponton, S., J. Dupouey, N. Brda, and E. Dreyer.
Annu. Conf. on Hydroponics. Hydroponic Soc. Nutr. 24:753765. 2002. Comparison of water-use efficiency of
Amer., San Ramon, Calif. Lang, H.J. 1996. Growing media testing and interpre- seedlings from two sympatric oak species:
Caviglia. O.P. and V.O. Sadras. 2001. Effect of ni- tation, p. 123139. In: D.W. Reed (ed.). Water, genotype environment interactions. Tree
trogen supply on crop conductance, water- and media , and nutrition for greenhouse crops. Ball Physiol. 22:413422.
radiation-use efficiency of wheat. Field Crops Publ., Batavia Ill. Uva, W.L., T.C. Weiler, and R.A. Milligan. 1998.
Res. 69:259266. Larouche, R., A. Gosselin, and L.P. Vezina. 1989. A survey on the planning and adoption of zero
Elliot, G. 1990. Reduce water and fertilizer with Nitrogen concentration and photosynthetic runoff subirrigation systems in greenhouse
ebb-and-flow. Greenhouse Grower 8:7072, photon flux in greenhouse tomato production: operations. HortScience 34:660663.
7475. I. Growth and development. J. Amer. Soc. Hort. van Iersel, M.W. 1996. Improving water and fertil-
Gislerrd, H.R. and L.M. Mortensen. 1990. Relative Sci. 114:458461. izer efficiency in greenhouses. Ga. Floricult.
humidity and nutrient concentration affect nutri- Le Roux, X., T. Bariac, H. Sinoquet, B. Genty, C. 6:2223.
ent uptake and growth of Begonia hiemalis. Piel, A. Marriotti, C. Girardin, and P. Richard. van Iersel, M.W. and B. Bugbee. 2000. A semi-con-
HortScience 25:524526. 2001. Spatial distribution of leaf water-use tinuous, multiple chamber crop CO2 exchange
Israeli, Y., A. Schwartz, Z. Plaut, and D. Yakir. 1996. efficiency and carbon isotope discrimination system: Design, calibration and data interpreta-
Effects of light regime on 13C, photosynthesis within an isolated tree crown. Plant Cell Environ. tion. J. Amer. Soc. Hort. Sci. 125:8692.
and yield of field-grown banana (Musa sp., 24:10211032. Vandana, B.R.K. 1999. Physiological changes in
Musaceae). Plant Cell Environ. 19:225230. Mills, H.A. and J.B. Jones. 1996. Plant analysis sesbania species to reducing light intensities. J.
James, E.C. and M.W. van Iersel. 2001. Fertilizer handbook II. MicroMacro Publ., Athens, Ga. Agron. Crop Sci. 182: 4347.
concentration affects growth and flowering of Morvant, J.K., J.M. Dole, and E. Ellen. 1997. Irriga- Wright, R.D. 1986. The pour-through nutrient extrac-
subirrigated petunias and begonias. HortScience tion systems alters distribution of roots, soluble tion procedure. HortScience 21:227229.
36:4044. salts, nitrogen and pH in the root medium. Hort- Yelanich, M.V. and J.A. Biernbaum. 1990. Effect of
Klring, H.P. and W. Cierpinski. 1998. Control of Technology 7:156160. fertilizer concentration and method of application
nutrient solution concentration depending on Nemali, K.S. and M.W. van Iersel. 2004. Light on media nutrient content, nitrogen runoff and
greenhouse climate in a sweet pepper crop. Acta intensity and fertilizer concentration: II. Opti- growth of Euphorbia pulcherrima V-14 glory.
Hort. 458:141146. mal fertilizer solution concentration for species Acta Hort. 272:185189.

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