J. AMER. SOC. HORT. SCI. 128(3):441–446. 2003.

Boron Improves Growth, Yield, Quality, and

Nutrient Content of Tomato
Jeanine M. Davis,1 Douglas C. Sanders,2 Paul V. Nelson,2 Laura Lengnick,3 and Wade J. Sperry4
Department of Horticultural Science, North Carolina State University, Raleigh, NC 27695-7609
ADDITIONAL INDEX WORDS. Lycopersicon esculetum, potassium, calcium, hydroponics, fruit crack, shelf life
ABSTRACT. Boron deficiency in fresh-market tomatoes (Lycopersicon esculentum Mill.) is a widespread problem that
reduces yield and fruit quality but is often not recognized by growers. Tomatoes were grown in field and hydroponic
culture to compare the effects of foliar and soil applied B on plant growth, fruit yield, fruit quality, and tissue nutrient
levels. Regardless of application method, B was associated with increased tomato growth and the concentration of K,
Ca, and B in plant tissue. Boron application was associated with increased N uptake by tomato in field culture, but
not under hydroponic culture. In field culture, foliar and/or soil applied B similarly increased fresh-market tomato
plant and root dry weight, uptake, and tissue concentrations of N, Ca, K, and B, and improved fruit set, total yields,
marketable yields, fruit shelf life, and fruit firmness. The similar growth and yield responses of tomato to foliar and
root B application suggests that B is translocated in the phloem in tomatoes. Fruit from plants receiving foliar or root
applied B contained more B, and K than fruit from plants not receiving B, indicating that B was translocated from
leaves to fruit and is an important factor in the management of K nutrition in tomato.
Boron deficiency in fresh-market tomatoes (Lycopersicon foliar spray regime ensures a steady supply of B throughout the
esculentum Mill.) is often not recognized by growers. Boron growing season without the management concerns of supplying
deficiency, however, is widespread (Gupta et al., 1985) and can B as a dry formulation. Application through the drip-irrigation
cause serious yield reductions and uneven ripening of tomato system places B directly in the root zone and foliar B applica-
fruit (Adams, 1978). Boron becomes less available to plants as tions allow uptake directly through the leaves. The objective of
soil pH increases (Bunt, 1956). Therefore the practice of apply- this study was to determine if B, applied in a liquid formulation
ing lime to improve the uptake of other important nutrients can to the foliage and/or roots, would improve yields, fruit quality,
cause B deficiency (Fleming, 1980). Boron deficiency can also and nutrient content of tomatoes.
occur readily in tomatoes grown in areas with acid sandy soils
and heavy rainfall (Adams, 1986) because it is easily leached Materials and Methods
under such conditions (Mengel and Kirkby, 1987). These condi-
tions are common in the southeastern United States (Mortvedt HYDROPONIC STUDIES. Production of tomato seedlings for the
and Woodruff, 1993; Offiah and Axley, 1993) and increase the hydroponic study was carried out by seeding fresh-market ‘Ce-
difficulty of managing B nutrition in tomatoes in this region. lebrityʼ tomato in flats containing steam sterilized river bottom
Maintenance of high foliage K levels during tomato fruit devel- sand. Flats were placed in a deionized water intermittent mist
opment is recommended but is often difficult to achieve (Adams, bed under cycles of 23 °C day/18 °C night with an 18-h photo-
1986; Bradley and Flemming, 1960). Fruit become large sinks for period provided by cool-white fluorescent lamps. Eighteen days
K as they develop, thereby diminishing K levels in leaves that are following sowing, seedlings were transplanted to acid washed
needed for continued plant growth and physiological processes 3.75-L plastic containers containing continuously aerated nutri-
(Ho and Hewitt, 1986). Adequate B levels help to maintain leaf ent solutions and placed in a greenhouse. To eliminate light, the
K levels in tomato during fruit development (Sperry, 1995). plastic containers were sprayed with aluminum colored paint and
Blevins et al. (1993) reported that B has a major influence on polystyrene covers were made to fit tightly in the openings. Three
the plasma membrane of plant cells and ion transport and that B small holes were bored in each cover to accommodate two plants
amendments increased K, Ca, and Mg levels in soybean [Glycine (one plant per hole) and an air tube. A closed-circle air manifold
max (L.) Merr.] leaves. was constructed with Tygon tubing and attached to an air pump
Fresh-market tomato production commonly involves use of (Rev-A, Thomas Industries, Inc., Pittsburgh, Pa.) and acid washed
raised beds with black polyethylene mulch and drip-irrigation. air stones were attached to the manifold, placed in the containers,
Soluble fertilizers are often applied directly to the root zone and suspended 2.5 cm from the bottom of the container. Constant
through the drip-irrigation system. Although B can be supplied levels of a modified Hoagland solution minus the B (Maynard
to tomatoes as dry fertilizer (Mortvedt and Woodruff, 1993), dry and Hochmuth, 1997) were maintained in the plastic containers
B formulations are difficult to manage because typically very by adding double-deionized water daily. Nutrient solutions were
small amounts are required, it is subject to loss by leaching, and completely changed every 7 d.
the range of deficient to toxic levels of soil B is very narrow Three treatments were applied in this study: no B; 1 mg·L–1 B
(Reisenauer et al., 1973). as H3BO3 supplied in the nutrient solution; and a foliar application
Application of B to tomatoes via drip-irrigation or in a regular of 1.87 mg·L–1 B as N-Boron (4.5–0–0 with 3.3% B chelated with
mannitol) (Claw El, Divison of Brandt Consolidated, Pleasant
Received for publication 1 July 2002. Accepted for publication 14 Jan. 2003. Plains, Ill.) Treatments were started when seedlings were placed
1Associate professor, Mountain Horticultural Crops Research and Extension
in the 3.75-L containers. The B treatments were supplied con-
Center, 455 Research Drive, Fletcher, NC 28732.
2Professor.
tinuously in the nutrient solution and the foliar applications were
3 Consultant. applied every seven days. The foliar treatments were applied with
4Former graduate research assistant. a hand held spray bottle and sprayed until just before run off. To

J. AMER. SOC. HORT. SCI. 128(3):441–446. 2003. 441

prevent contamination of plant roots and growth solution with Lateral shoots, 5 to 10 cm long, were removed from the base of
the foliar spray treatment, containers were taken to a separate the plant up to, but not including, the shoot immediately below
room before treatment and the containers and plants up to the the first flower cluster. Recommended weed, disease, and insect
first leaves were protected by securing a 56.7-L plastic bag tightly management practices (North Carolina State Univ., 1993) were
around the stem. A new bag was used for each container and used throughout the course of the study.
each time a treatment was applied. Plants were harvested 56 d Treatments were assigned to the experimental plots as a ran-
after transplanting. To remove B treatment surface residues from domized complete block design with three treatments and five
roots and shoots, the tissues were rinsed under warm tap water replications. The three treatments consisted of no B fertilization,
for 15 s, washed for 30 s in 0.2 N HCl, rinsed under warm tap a weekly foliar B application, and a weekly soil B application.
water for 15 s, and rinsed for 30 s in distilled water (Williams and Foliar B was applied at 0.56 kg·ha–1 B as N-Boron using a back-
Nelson, 1992). All samples were weighed, dried at 70 °C for 72 pack sprayer. Weekly soil B was applied at 0.56 kg·ha–1 through
h, reweighed, ground in a stainless steel Wiley mill) to a particle a line of drip-tape separate from the irrigation drip-tape, placed
size ≤1 mm (20-mesh screen), and dry-ashed at 500 °C. Samples directly under the polyethylene mulch and on top of the soil. The
were analyzed for total N by a semimicro-Kheldahl procedure in separate line could be attached or unattached to the irrigation drip-
which salicylic acid was added as a pretreatment to digestion to tape system when soil B treatments were applied. A polyvinyl
aid in the reduction of NO3 (Eastin, 1978). Boron was determined chloride pipe injection manifold with a backflow prevention valve
by the curcumin procedure (Grinstead and Snider, 1967) and was constructed (Lancaster et al., 1998), connected to the soil B
K, Ca, Mg, and S (sulfur) were analyzed by atomic absorption treatment drip system, and used to apply the soil B treatments.
spectroscopy (Christian and Feldman, 1970). Phosphorus was Application of the experimental treatments began at first anthesis
analyzed colorometrically (Chapman and Pratt, 1961) using a and continued for 8 weeks for a total of 4.48 kg·ha–1 B. Leaflet
UV/VIS spectrophotometer (Perkin-Elmer, Norwalk, Conn.). An samples were taken from the petiole of the fourth leaf from the
estimate of total nutrient uptake was calculated by multiplying growing tip to evaluate B effects on leaflet K and other nutrient
the concentration of nutrient (mg nutrient/g plant tissue) by the changes from first anthesis through fruit development. Leaflets
total dry weight of the specific plant tissue. were sampled 7 d following the first B applications and weekly
This study was repeated three times using a randomized com- thereafter for 8 weeks. Percent fruit set was evaluated by tagging
plete block design with three treatments and five replications. A newly opened blossoms once a week and counting the number
fourth study was conducted that included the three treatments of tagged blossoms which set fruit 1 week later. Plant height was
described previously, plus an additional foliar + root B applica- measured from mulch surface to the growing tip at time of last
tion treatment. The foliar+root B application treatment supplied 1 harvest. Fruit were harvested six times at the breaker stage (just
mg·L–1 B from H3BO3 in the nutrient solution and 1.87 mg·L–1 B as showing pink at the blossom end), graded, and weighed weekly.
an N-Boron foliar spray. Except for the addition of the foliar+root Fruit were graded into U.S. Combination, U.S. no. 3, and culls
B application treatment, the fourth study was conducted using the (U.S. Dept. of Agriculture, 1961). Yields reported are marketable
same materials and methods as described previously. fruit (U.S. Combination and U.S. no. 3) or total yield (marketable
The results of all four experiments were subjected to ANOVA. plus culls). Fruit were culled for cracks, zippers, cat-facing, and
No treatment by experiment interactions were found, so the data other damage. All cracked fruit were weighed.
were grouped as one experiment with four treatments and unequal Forty randomly selected fruit per plot were placed on tables
replication (n = 5 for the foliar + root treatment, n = 20 for all in a well ventilated laboratory at 22 °C and evaluated for shelf
other treatments). The grouped data were subjected to analysis life. Fruit were placed on paper and arranged so they were not
of variance and, where appropriate, to the F protected LSD mean touching. Fruit were checked daily and removed at first sign of
separation test using SAS (SAS Institute, Cary, N.C.). breakdown due to dehydration or disease and days to removal
FIELD STUDY. This field study was conducted on a Norfolk loamy were recorded.
sand (fine loamy, siliceous, thermic Typic Paleudult) in Clayton, Another randomly selected group of fruit was used for nutri-
N.C. Soil had a pH of 6.6 and contained less than 0.1 mg·kg-1 B. ent analysis. Boron treatment surface residues were removed
This site was selected because B deficiency symptoms are likely from this fruit using the method of Williams and Nelson (1992)
to occur in tomato plants when soil B concentrations are below a described previously. Fruit were cut and separated into 600 g
critical level of 0.1-0.7 mg·kg–1 (Maynard and Hochmuth, 1997). allotments per plot, then dried at 70 °C for 96 h, weighed for
Soil B levels were analyzed colorometrically using the hot-water- dry matter, ground, and analyzed for tissue nutrient content as
soluble B extraction method of McGeehan et al. (1989). described previously.
The experimental site was prepared for tomato production An additional group of randomly selected fruit was harvested
by the application of 11.2 kg·ha–1 P, 84 kg·ha-1 K, and 56 kg·ha-1 from each plot at the breaker stage for crack-point testing. Forty
N broadcast applied and incorporated into the soil. Beds were fruit per plot were placed on paper, and arranged so they were
formed and fumigated with a mixture of 67% methylbromide not touching, on tables in a well-ventilated laboratory with day/
and 33% chloropicrin (CC13NO2) at 224 kg·ha–1 4 weeks before night cycles of ≈12 h at 22 to 23 °C. The fruit were allowed to
planting. At the same time, black polyethylene mulch (1.5 m attain uniform ripeness of the light red stage and then subjected
wide × 0.04 mm thick) and drip-tape (61 cm emitter spacing) to crack point testing using an Instron Universal Tester (model
(Typhoon: Netafim, Valley Stream, N.Y.) were applied. The TM; Instron Corp., Atlanta, Ga.). Pressure was exerted midway
drip-tape was placed 4 cm deep and 10 cm from the center of between the blossom and stem ends of the fruit with a 10.16 cm2
the bed. Plots consisted of four 4.6 m long rows spaced 1.5 m plate moving at constant speed. The force needed to cause the
apart with the outer two rows serving as guard rows. Six-week- skin to crack was recorded.
old greenhouse-grown ‘Celebrityʼ transplants were mechanically Following final harvest, two plants per plot were harvested
set 46 cm apart in-row on 3 May 1994. Plants were staked and and prepared for tissue analysis as described previously. Samples
trained to the string weave system (Konsler and Gardner, 1990). were analyzed for total N, P, K, B, Ca, Mg, and S as described

442 J. AMER. SOC. HORT. SCI. 128(3):441–446. 2003.

previously. An estimate of total nutrient uptake was calculated translocated from foliage to roots. Roots from the control plants,
by multiplying the concentration of nutrient (mg nutrient per g although clearly B deficient, still contained small amounts of B
plant tissue) by the total dry weight of the specific plant tissue (Table 1). We analyzed seed from ‘Celebrityʼ tomato and 20-d-
or organ. old ‘Celebrityʼ seedlings and found an average of 27 mg·kg–1 B
Data were subjected to analysis of variance and, where ap- in seed and 10 mg·kg–1 B in seedlings.
propriate, to the F protected LSD mean separation test using SAS Historically, B has been considered to be a phloem immobile
(SAS Institute, Cary, N.C.). element (Oertli and Richardson, 1970). One of the exceptions
appears to be some plants that translocate sugar alcohols such as
Results and Discussion mannitol and sorbitol. Sugar alcohols, however, are not present in
tomatoes (Brown et al., 1999). The apparent phloem transport of
Hydroponic studies. Plants not treated with B had lower shoot B that was observed in this study has been reported by other re-
and root dry weights than plants treated with a B application (Table searchers working with tomatoes (Cerda and Roorda van Eysinga,
1). Stunted growth in B deficient conditions has been observed 1981) as well as a variety of other crops: in B deficient broccoli
in tomato and several other crops (Gupta and Cutcliffe, 1985; (Brassica oleracea L.) (Benson et al., 1961; Shelp, 1987) and
Nelson et al., 1977). Shoot N, Mg, and S concentrations were grape (Vitis vinifera) (Scott and Schrader, 1947), or following
not affected by B treatments (data not shown); however, plants foliar applications of B on apple [Malus ×sylvestris (L.) Mill. var.
treated with root and root+foliar applied B had higher shoot P, domestica (Borkh.) Mansf.], pear (Pyrus communis L.), prune
K, and Ca levels than plants not treated with B (Table 1). Boron (Prunus domestica L.), and cherry (P. avium L.), (Picchioni et
applied to foliage and/or roots improved uptake of N, K, Ca, and al., 1995). In a study of the mobility of foliar-applied labelled
B into shoots, roots, and fruit (Table 2). Boron concentration was 10B in peach (Prunus persica), Shu, et al. (1993) showed that B

lowest in shoots of plants receiving no B (Table 1). Potassium is swiftly (within 72 h of foliar application) re-translocated in the
concentrations in tomato shoots responded positively to B sup- phloem in priority to sinks throughout the plant, regardless of the
plied to the roots or as a foliar amendment, which agrees with distance from the source application. These workers conclude that
the work of Cerda and Roorda van Eysinga (1981) on tomato and the commonly held view of B as a phloem immobile element may
Schon and Blevins (1990) on soybean. be partly due to the difficulty of using conventional non-tracer
Roots from plants not treated with B had the lowest Ca and methods to observe the movement of the small amounts of B
B concentration (Table 1) and were stunted with blackened typically involved in plant growth and development.
tips. These are classic symptoms of B deficiency (Maynard and Field study. PLANT GROWTH AND YIELD. Tomato response to B
Hochmuth, 1997). Roots from plants receiving B had none of application under field culture supported the results observed in
these symptoms. Lack of B deficiency symptoms and high tis- the hydroponic experiments. Tomatoes receiving soil or foliar-
sue B concentration (Table 1) and uptake (Table 2) in roots from applied B had better growth and were more productive compared
plants that received only foliar applied B suggests that B was to plants that did not receive B (Table 3). Fruit set occurred more
Table 1. Shoot and root tissue nutrient concentration and dry matter of 56-d-old ‘Celebrityʼ tomato grown in hydroponic culture and treated with
foliar, root, foliar+root, or no applied boron.
Nutrient concnz Dry
Boron P K Ca B matter
treatment mg·g-1 mg·kg-1 (g)
Shoot
None 7.0 by 47.4 c 16.4 c 16.6 c 2.35 b
Foliar applied 6.7 c 53.0 b 17.9 b 95.6 a 3.11 a
Root applied 7.2 a 62.1 a 18.8 a 73.7 b 3.35 a
Root+foliar 7.2 a 62.8 a 18.9 a 94.0 a 3.16 a
Root
None 8.3a 46.1 a 6.3 b 20.4 b 0.64 b
Foliar applied 8.5 a 45.4 a 8.1 a 68.0 a 1.06 a
Root applied 8.8 a 45.4 a 8.0 a 71.7 a 1.16a
Root+foliar 8.5 a 45.6 a 8.0 a 69.4 a 1.05 a
Reported tissue concentration and dry matter means are the combined treatment means of all experiments.
z

Mean separation within columns by F protected LSD at P ≥ 0.05.

y

Table 2. Nutrient uptake response to foliar, root, or no applied B in tomatoes under hydroponic culture.
Total uptake (shoot + root) Shoot content Fruit content
Boron N K Ca B N K Ca B N K Ca B
treatmentz g/plant mg/plant g/plant mg/plant g/plant mg/plant
None 0.17 by 0.11 b 0.030 b 0.03 b 0.14 b 0.09 b 0.030 b 0.02 c 0.03 b 0.02 b 0.003 b 0.006 b
Foliar applied 0.30 a 0.23 a 0.070 a 0.40 a 0.23 a 0.18 a 0.060 a 0.29 a 0.07 a 0.05 a 0.010 a 0.100 a
Soil applied 0.29 a 0.25 a 0.070 a 0.36 a 0.22 a 0.20 a 0.060 a 0.26 b 0.07 a 0.05 a 0.008 a 0.090 a
Soil+foliar 0.29 a 0.25 a 0.070 a 0.40 a 0.23 a 0.20 a 0.060 a 0.30 a 0.06 a 0.05 a 0.008 a 0.100 a
Reported nutrient uptake means are the combined treatment means of all experiments.
z

Mean separation within columns by F protected LSD at P ≥ 0.05.

y

J. AMER. SOC. HORT. SCI. 128(3):441–446. 2003. 443

Table 3. Influence of foliar and soil applied boron on field-grown tomato quality and yield parameters.
Plant growth and fruit yield Fruit quality
Plant Plant Fruit Total Marketable Shelf Crack Concentric Radial
Boron dry wt ht set yield yield Jumbo Extra large Large life point crack crack
treatment (g) (cm) (%) (Mg.ha–1) (%) (Mg·ha–1) (d) (N) (Mg·ha–1)
None 405 b 122 b 56.1 b 37.0 b 82.5 b 2.66 b 4.75 b 10.8 b 8.0 b 47.0 b 1.29 a 0.43 a
Foliar 457 az 135a 80.4 a 49.6 a 87.5 a 4.69 a 8.96 a 14.4 a 13.0 a 53.2 a 0.78 b 0.15 b
Soil 454 a 137 a 73.2 a 48.1 a 87.7 a 4.46 a 8.67 a 14.0 a 14.2 a 55.4 a 0.79 b 0.26 ab
zMean separation within columns by F protected LSD at P ≥ 0.05.

Table 4. Shoot and fruit tissue nutrient concentration of fresh-market cracks than plants not treated with B (Table 3). These results
field-grown tomato treated with foliar and root applied boron. suggest that B may be important in reducing the incidence of fruit
Nutrient concnz cracking. Dixon et al. (1973) reported that cracking in apple was
Boron K Ca B entirely eliminated by sprays containing B. This may be due to
treatment (g·kg-1) (mg·kg-1) the effects of B on membranes and cell walls. Many studies have
Shoot shown that B helps maintain membrane stability (Ginzburg, 1961;
None 29.8 b 22.9 b 29.7 b Pilbeam and Kirkby, 1983; Yamouchi et al., 1986). Yamouchi et
Foliar applied 48.7 a 24.8 ab 80.3 a al. (1986) reported that B deficiency in tomato actually induced
Root applied 54.8 a 25.1 a 78.2 a a reduction in the amount of Ca associated with the pectin frac-
Fruit tion of tomato leaf cell walls. They found 67% of the total B in
None 35.2 b 2.4 a 17.1 b the cell wall fraction of tomato leaf tissues and suggested that
Foliar applied 46.9 a 2.6 a 27.3 a B may function in cell wall metabolism by maintaining the Ca-
Root applied 44.0 a 2.6 a 28.8 a pectin association.
TISSUE NUTRIENT CONTENT. As in the hydroponic study, K con-
zMean separation within columns by F protected LSD at P ≥ 0.05. centration (Table 4) and uptake (Table 5) were highest in shoot
tissues when plants were treated with foliar or soil B. Calcium
than 70% of the time on plants receiving foliar or soil applied B and B concentration (Table 4) and uptake (Table 5) were lowest
compared to only 56% of the time on plants that did not receive in plant tissues when plants were not treated with B, while Ca
B (Table 3). Plants receiving foliar or soil B had higher total and was highest in plants treated with soil B. This agrees with stud-
marketable yields than plants receiving no B (Table 3). ies by Hill and Grant (1935), and Minarik and Shive (1939) who
These results agree with other studies in which soil applied B demonstrated that adding B to the growing medium increased
(Gulati et al., 1980) or foliar applied B (Gascho, 1993; Oplinger et the amount of Ca in plants. It is interesting that although total
al., 1993; Schon and Blevins, 1990; Weaver et al., 1985) improved plant uptake of N, K, and Ca increased with B application (Table
yields in tomato, soybean, and snap bean (Phaseolus vulgaris 5), the concentration of N did not (data not shown) indicating
L. var. vulgaris). De Magalhaes et al. (1981) reported increased that N uptake keeps pace with growth while increased K and Ca
tomato fruit yields and quality with soil applied boron but had no uptake exceeded the increased growth resulting from B applica-
response to foliar applied boron. Oplinger et al., (1993) observed tions. Shoot N, P, Mg, and S concentrations were not altered by
that B applied to soybean foliage at initial flowering increased B treatment (data not shown).
yields by 3% and Gascho (1993) reported yield increases when Widders and Lorenz (1982) demonstrated that redistribution
B and N were applied to soybean foliage during reproductive of K from vegetative tissues contributed up to 12% of the total K
development. Yields of jumbo, extra large, and large fruit were content of tomato fruit with subsequent foliar K levels decreasing
greater when plants were treated with foliar or soil applied B than during fruiting by 20% to 40%. We wanted to observe whether
if plants received no B (Table 3). Yields of medium and small soil or foliar B amendments would aid in maintenance of foliar K
fruit were not affected by B (data not shown). concentrations during fruit development. There was a significant
FRUIT QUALITY. Foliar and soil applied B increased shelf life and B × week interaction (Fig. 1). As fruit development progressed,
fruit crack point compared to nontreated plants (Table 3). In this leaflet K levels declined from >4% to <2% during weeks 3 through
study, plants treated with soil applied B had a higher concentration 8 in plants not treated with B. Potassium levels in plants treated
of Ca than plants not treated with B, but fruit Ca concentrations with soil or foliar B declined from ≈4.5% at week 2 to just over
were not significantly different (Table 4). Similar results were 3% at week 6 and maintained these levels through week 8. Based
obtained in a controlled environment study (Sperry, 1995). on these results, applied B appears to be important in maintaining
Concentric and radial fruit crack are two common types of high leaflet K concentrations in field grown tomato, especially
cracks which occur on tomatoes (Peet, 1992; Sperry, 1995). Plants after 5 to 6 weeks of fruit development. Similar results were
treated with foliar B had less fruit with concentric and radial found in a 2-year field study by Sperry (1995). No significant
Table 5. Total nutrient uptake response to foliar or root applied B in field-grown tomatoes.
Boron Total uptake (shoot + root) (kg·ha–1) Shoot uptake (kg·ha–1) Root uptake (kg·ha–1)
treatment N K Ca B N K Ca B N K Ca B
None 397 bz 233 b 137 b 0.22 b 329 b 174 b 133 b 0.17 c 67 b 60 b 4.0 b 0.05 b
Foliar applied 475 a 427a 169 a 0.71 a 386 a 319 a 163 a 0.53 a 89 a 108 a 6.0 a 0.18 a
Soil applied 466 a 456 a 169 a 0.68 a 382 a 358 a 163 a 0.51 b 84 a 98 a 6.0 a 0.17 a
zMean separation within columns by F protected LSD at P ≥ 0.05.

444 J. AMER. SOC. HORT. SCI. 128(3):441–446. 2003.

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