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Nutrient uptake and yield of sweet pepper as affected by stage of development

and N form

Article  in  Journal of Plant Nutrition · November 1991

DOI: 10.1080/01904169109364275

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Héctor Marti Harry A. Mills

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 Journal of Plant Nutrition

ISSN: 0190-4167 (Print) 1532-4087 (Online) Journal homepage: http://www.tandfonline.com/loi/lpla20

Nutrient uptake and yield of sweet pepper as

affected by stage of development and N form

Hector R. Marti & Harry A. Mills

To cite this article: Hector R. Marti & Harry A. Mills (1991) Nutrient uptake and yield of sweet
pepper as affected by stage of development and N form, Journal of Plant Nutrition, 14:11,
1165-1175, DOI: 10.1080/01904169109364275

To link to this article: http://dx.doi.org/10.1080/01904169109364275

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 JOURNAL OF PLANT NUTRITION, 14(11), 1165-1175 (1991)

NUTRIENT UPTAKE AND YIELD OF SWEET PEPPER AS AFFECTED

BY STAGE OF DEVELOPMENT AND N FORM

Hector R. Marti and Harry A. Mills

University of Georgia, Department of Horticulture, Athens, GA 30602

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ABSTRACT: Uptake of NO3, NH4+, P, K++, Ca++ and Mg++, as influenced by the
stage of plant development and three NO3-:NH4+ ratios (1:0, 1:1, and 0:1), was
determined for sweet pepper (Capsicum annuum L. cv. 'California Wonder').
Uptake was highest during fruit development and immediately after fruit harvest,
indicating that fruit removal promotes nutrient uptake. When NO3- and NH4+ were
supplied in equal concentrations, NO3- was absorbed more readily. Each
increment in NH4+ + ++
decreased the uptake of K , Ca , and Mg ++
by fruit tissue,
while no significant effect on the N and P content of the fruit was observed.
Ammonium nutrition reduced plant dry weight and fruit yield in comparison to
NO3-. Results from this study suggest that NO3- is the preferred N form, and that
fertilizer application should be scheduled according to specific physiological stages
to maximize nutrient uptake. Nutrient content of vegetative tissue was not
indicative of potential yield.

INTRODUCTION
Correlating nutrient uptake by a crop to specific physiological stages of plant
development is essential to developing a fertility program that maximizes nutrient
uptake and accumulation to prevent periodic nutrient stress during the growth
cycle. Studies with various crops (8,11,13) show different demand periods for
the uptake of NO3' and NH4+. Previous studies with sweet pepper (10,14)

1165

Copyright © 1991 by Marcel Dekker, Inc.

 1166 MARTI AND MILLS

emphasized the distribution and accumulation of nutrients in the vegetative tissue

at different stages of development rather than uptake or demand periods for
nutrient acquisition. The impact of N form on nutrient uptake has not been
evaluated for sweet pepper, although studies with other crops have shown a
significant effect of N form on nutrient uptake. With lima beans (8) and
southempeas (13), ammonium reduced dry weight and total nutrient uptake but not
the demand periods for nutrient uptake. Ammonium enhanced the vegetative
growth of tomatoes (3), but reduced fruit weight (3,12). Ammonium and urea are
extensively used in the sweet pepper industry, even though NH4+ nutrition can
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reduce the dry weight of sweet pepper seedlings (5), fruit yield (10,18), and
nutrient accumulation in vegetative tissue (4,5,18). The objectives of this study
were to determine the influence of stage of plant development and N form on
nutrient uptake and yield of sweet pepper.

MATERIALS AND METHODS

Bell pepper (Capsicum annuum L. cv. 'California Wonder') were cultured in
15 liter pots containing a modified Hoagland's solution with elemental concen-
trations (mg/1) as follows: N (150), P (15), K (150), Ca (200), Mg (30), B (0.25),
Cu (0.15), Fe (2.5), Mn (0.25), Mo (0.10), and Zn (0.25), under greenhouse con-
ditions (day/night temperature 32/21°C, photoperiod 13.5 hr). Three NO3" : NH4+
ratios were tested: 1:0, 1:1, and 0:1. Calcium nitrate was the N0 3 'N source, and
(NH^SO,, the NH4+ N source. CaCl2 was used to balance the Ca** concentration.
The N concentration was 75 mg/L the first week, and 150 mg/L the second week
until harvest. The concentrations of the other essential elements were the same
for all the treatments, with SO4= and Cl' being variable. The statistical design was
a completely randomized design with 4 single-plant replications. Transpired water
was replaced daily, and solutions were changed weekly. Nutrient uptake was
calculated at the end of each week as being the difference between initial and final
solution concentrations. Flowering occurred at week 4, with the first fruits
 NUTRIENT UPTAKE 1167

developing to half size at week 6. Two harvests of the fruits were made, one at
week 8 and the other at week 12. At both harvests, all fruits were removed when
the fruits of the first cluster were at the mature green stage (full size).
Plants were dried at 70° C, divided into roots, stems, leaves, and fruits, and
ground in a Wiley mill to pass through a 20-mesh screen. Total N was deter-
mined by Kjeldahl digestion, with an Autoanalyzer utilized to determine NH4+, and
NO3' in tissue and/or nutrient solution (6). Other elements were determined by
plasma emission spectrophotometry (7).
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The data were subjected to analysis of variance. Duncan's test was used to
compare uptake means at stages of development within each NO3' : NH4+ ratio.
Regression analysis was performed on data for NO3", NH4\ P, K, Ca, and Mg
composition, dry weight, and yield of plant parts.

RESULTS AND DISCUSSION

Stases of Development: In general, with the 1:0 and 1:1 NO3' : NH4+ treatments,
nutrient uptake increased from flowering until the fruits were one-half their mature
size, but decreased as fruits reached their full size (Figures 1 to 6), The removal
of all the fruits produced a sharp increase in nutrient uptake (Postharvest, Figure
1 to 6), which decreased again when the fruits of the most mature cluster reached
their final size.
Miller et al. (10) and Tapia and Dabed (14) reported that stage of pepper
development influenced nutrient accumulation in tissue. Thomas and Heilman
(15) and Batal and Smittle (1) reported an increase in N content of leaves after
harvest. In these previous studies direct uptake was not measured, and the N form
was not considered. Our results suggest that nutrient uptake is reduced as fruits
from the most mature cluster approach their final size. This reduced nutrient
uptake at fruit maturity is possibly due to a competition for energy between
nutrient uptake and fruit growth. Hall and Brady (2) suggested that increased
respiration of the fruit caused a drop in the net CO2 assimilation rate. Thus, fruit
 1168 MARTI AND MILLS

development is likely to affect the energy available for other plant processes,
including nutrient uptake. However, other mechanisms cannot be discarded, since
Ca**, which is taken up passively, showed the same pattern of uptake as other
elements taken up actively.
To maximize nutrient uptake, fertilizer applications should be scheduled
according to demand periods and actual uptake of each nutrient. Based on our
study, supplemental nutrient applications would be most effective if applied at the
initiation of the first fruits and continued through fruit development. Applications
of N, P, K+, Ca", and Mg++ should be supplemented after each fruit harvest.
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N form: Nutrient uptake declined with each increment of NH4+ with all stages
of development (Figure 1 to 6).
When NO3' and NH/ were at the same concentration (1:1 treatment), NO3"
uptake was higher than NH4+ uptake (Figure 1). In addition, NH/ uptake was
higher with the 1:1 treatment than with NH4+ as the sole source of N (Figure 2).
These results show a preference of the pepper plant for NO3' in respect to NH4+.
The decline in P and K+ uptake with NH/ was linear at most stages of
development, and the largest decline occurred generally with the 0:1 NO3' : NH4+
treatment (Figures 3 and 4).
For Ca" and Mg4*, the difference between the 1:1 and 0:1 NO3' : NH+ ratios
was not significative except at postharvest, and the linear and quadratic response
was the most common (Figures 5 and 6). This indicates that for these cations,
uptake is seriously reduced when NH/ is 50% or more of the N form.
The negative effects of NH4+ as the primary source of N are reflected in the
dry weight of plant parts and fruit yield. The dry weight of each plant part, and
both total and marketable yields, decreased with each increment of NH4+ in the N
form (Table 1). The same type of response was found by others for the dry
weight of pepper plants (5) and for fruit yield (9,18). Nitrogen significantly
influenced the final elemental composition of plant parts (Table 1). Kjeldahl N
in plant tissue increased with each increment of NH4+ in the N form. This was
 NUTRIENT UPTAKE 1169

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Downloaded by [. Hector Marti] at 06:39 21 April 2016

0.5
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de

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0.0
11 15OPI îmRV
STAGES OF DEVELOPMENT
SU
Figure 1. Nitrate uptake as affected by stage of development and N form. Small
letters indicate differences between stages of development within each N form
(Duncan, 5% level).

1.0
INQ3:NH4 IM N03-.NH4 OM L : LINERR
0.9 1 Z LEVEL
a

a
J3 0.7

D>0.6

ai 0.6
<r 0.4

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0.0
STAGES OF DEVELOPMENT
Figure 2. Ammonium uptake as affected by stage of development and N form.
Small letters indicate differences between stages of development within each N
form (Duncan, 5% level).
 1170 MARTI AND MILLS

0.3
BN03:NH4 1:0 i N03:NH4 1:1 !N03:rW4 0:1
a Ls LINEflR.
5S Qs QUñDRñTIC
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a
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o. a 1 1 I
t3>
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9
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Downloaded by [. Hector Marti] at 06:39 21 April 2016

•
d
bc
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0.0 ÍP-
_UUM
STRGES OF DEVB-OPMB-JT

Figure 3. Phosphorus uptake as affected by stage of development and N form.

Small letters indicate differences between stages of development within each N
form (Duncan, 5% level).

2. Ö
Z. 0 S N03:NH4 1:0 • NQ3:NH4 1 1 B ^03:^4 0:1
2. 4 L = LINEflR.
Q : SUñDRRTlC
Z. C
a a 1 X LEVEL
e.oi-
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— i. 8 a a
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4
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lb
^ o.
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o. 2
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M™ b b b b

3LuUM 1 ZFODT W 1/ KJLL h¿

STRGES 0 F DEVELOPMENT
Figure 4. Potassium uptake as affected by stage of development and N form.
Small letters indicate differences between stages of development within each N
form (Duncan, 5% level).
 NUTRIENT UPTAKE 1171

1.4
N03:MI4 hi i NÛ3:NH4 OM
1.2 a L- LIHEflR
Q: GUflDRnnc
1 X LEVEL
Downloaded by [. Hector Marti] at 06:39 21 April 2016

c b
0.0
TffTW LJTBZ
7T5T
STRGES OF DEVELOPMENT

Figure 5. Calcium uptake as affected by stage of development and N form.

Small letters indicate differences between stages of development within each N
form (Duncan, 5% level).

0.4

LINEflR
Qunonnnc
I X LEVEL

o.o -r
STRGES OF DEVELOPMENT

Figure 6. Magnesium uptake as affected by stage of development and N form.

Small letters indicate differences between stages of development within each N
form (Duncan, 5% level).
 1172 MARTI AND MILLS

of sweet pepper as influenced by three NOj-:NH4 ratios

Plant N N P K Ca" Mg Dry Wt
part ratio W (%) W m (*) (g) (kg/pl)
Total Mktbl

I1 1 1 1
1:0 4.6 5.5 0.6 33.80
Roots 1:1 5.8 0'.9 3.9 0Í6 0.2 21.42
0:1 5.7 3.5 1.2 5.6 0.2 3.80
1 1 1 1 1
0.4 4.5 0q6 63.47
Downloaded by [. Hector Marti] at 06:39 21 April 2016

1:0 2.3 1.6

Stems 1:1 3.8 0.5 3.4 1.3 0.5 34.00
0:1 6.5 0.8 2.5 1.2 1.0 2.05
ns 1 1 1 1
1:0 5.0 0.6 7.1 6.7 1.6 78.67
Leaves 1:1 6.6 0.7 7.3 3.2 0.6 55.25
0:1 6.4 0.9 4.3 2.8 0.6 4.77
ns ns 1 1 1 1
1:0 3.8 0.7 4.2 0.3 0.4 121.21 1.93 1..16
Fruits 1:1 4.3 0.7 3.7 0.2 0.3 106.88 1.50 0..63
0:1 5.0 0.4 1.7 0.1 0.1 4.27 0.66 0..00

Significance at the 1 Z level: l:linear, q:quadratic, ns:no

significant.

attributed to a increase in % N due in part to a decrease in dry weight and thus

a concentration effect. Therefore, measurement of Kjeldahl N in sweet pepper
plant parts would not be a useful indicator of the N status of the plant to predict
sufficiency for maximum yield. This is in agreement with the findings of Wilcox
et al. (16) and Hartman et al. (3) for tomato. Phosphorus in vegetative plant tissue
increased with each increment of NH/. This was possibly due to inorganic P
acting as counterion of NH/, as suggested by others (5,16). In the fruits, the
differences were not significant. Potassium in all plant parts decreased linearly
with each increment of NH4+. Ikeda and Osawa (4) reported that NH/ decreased
the K+ content of pepper seedlings when compared to NO3\ Calcium content of
stems, leaves, and fruits showed the same response as K+—a linearly decline with
 NUTRIENT UPTAKE 1173

each increment of NH/. In the roots, there was a sharp increase in Ca** with the
high NH,+ ratio (0:1). Probably the poor root growth with NH4+ resulted in Ca**
being concentrated in root tissue. Magnesium in the various plant parts generally
decreased as NH/ increased. This response is in agreement with the findings of
Ikeda and Osawa (4) for leaves of sweet pepper seedlings.
Despite the negative effects of NH4+ on plant growth, all leaf nutrient
concentrations were above the levels given by Winsor and Adams as sufficient
(17). These results indicate that the nutrient concentration of pepper leaves is not
Downloaded by [. Hector Marti] at 06:39 21 April 2016

a good indicator of potential yield unless the concentration is related to N form

and growth parameters.
In conclusion, nutrient uptake was influenced by physiological stage and N
form. Generally, nutrient uptake decreased as fruits reached full size, but
increased with fruit removal. In order to maximize nutrient uptake and yield,
supplemental fertilizer should be applied prior to fruit expansion and continued
through fruit development. Since selectivity for NO3" over NH4+ was observed,
NO3" should be at least 50% of the N form supplied to peppers through fertilizer
applications.

REFERENCES:
1. Batal, K.M., and D.A. Smittle. 1981. Response of bell pepper to irrigation,
nitrogen, and plant population. J. Amer. Soc. Hort. Sci. 106(3):259-262.

2. Hall, A.J., and C.D. Brady. 1977. Assimilate source-sink relationships in

Capsicum annuum L. II. Effects of fruiting and defloration on the
photosynthetic capacity and senescence of leaves. Aust. J. Plant Physiol.
54:623-636.

3. Hartman, P.L., H.A. Mills, and J.B. Jones, Jr. 1986. The influence of nitrate:
Ammonium ratios on growth, fruit development, and element concentration
of 'Floradel' tomato plants. J. Amer. Soc. Hort. Sci. 111(4):487-490.

4. Ikeda, I., and T. Osawa. 1979. Comparison of adaptability to nitrogen source

among vegetable crops. I. Growth response and nitrogen assimilation of fruit
vegetables cultured in nutrient solution containing nitrate, ammonium, and
nitrite nitrogen. J. Jap. Soc. Hort. Sci. 47:454-462.
 !
1174 MARTI AND MILLS

5. Ikeda, I., and T. Osawa. 1983. Effects of ratios NO3- to NH4+ and concen-
tration of each N source in the nutrient solution on growth and leaf N
constituents of vegetable crops and solution pH. J. Jap. Soc. Hort. Sci.
52(2):159-166.

6. Isaac, R.A. and W.C. Johnson. 1976. Determination of total nitrogen in plant
tissue. J. Assn. Official Anal. Chem. 59:98-100.

7. Jones, J.B., Jr. 1977. Elemental analysis of soil extracts and plant tissue ash
by plasma emission spectroscopy. Comm. Soil Sci. Plant Anal. 8:349-365.

8. McElhannon, W.S., and H.A. Mills. 1978. Influence of percent NO3-/NH4+

Downloaded by [. Hector Marti] at 06:39 21 April 2016

on growth, N absorption, and assimilation by lima beans in solution culture.

Agron. J. 70:1027-1032.

9. Milhet, Y., and Costes C. 1986. Influence de la forme et de la dose d'azote

de la fumure minerale sur la production de fruits de plusieurs plantes
maraicheres suivant les teneurs de l'air en dioxyde de carbone. Hort. Abs.
59(9):749.

10. Miller, C.H., R.E. McCollum, and S. Claimon. 1979. Relationships between
growth of bell peppers (Capsicum annuum L.) and nutrient accumulation
during ontogeny in field environments. J. Amer. Soc. Hort. Sci. 104:852-
857.
11. Mills, H.A., and W.S. McElhannon. 1982. Nitrogen uptake by sweet corn.
HortScience 17(5): 743-744.

12. Pill, W.G., and V.N. Lambeth. 1980. Effect of soil water regime and
nitrogen form on blossom-end rot, yield, water relations, and elemental
composition of tomato. J. Amer. Soc. Hort. Sci. 105(5):730-734.

13. Sasseville, D.N., and H.A. Mills. 1979. N form and concentration: Effects
on N absorption, growth, and total N accumulation with southernpeas. J.
Amer. Soc. Hort. Sci. 104(5):586-591.

14. Tapia, M.L., and R. Dabed. 1984. Nutrient uptake by sweet pepper brown
in quartz, pp. 683-696. Proc. Sixth. Congress Soilless Culture.

15. Thomas, J.R., and M.D. Heilman. 1964. Nitrogen and phosphorus content
of leaf tissue in relation to sweet pepper yields. Proc. Amer. Soc. Hort. Sci.
85:419-425.
 NUTRIENT UPTAKE 1175

16. Wilcox, G.E., C.A. Mitchel, and J.E. Hoff. 1977. Influence of nitrogen form
on exudation rate, and ammonium, amide, and cation composition of xylem
exudate in tomato. J. Amer. Soc. Hort. Sci. 102:192-196.

17. Winsor, G., and Adams P. 1987. Diagnosis of mineral disorders in plants.
Vol. 3: Glasshouse crops. Chptr 4: Peppers. Chemical Publishing Co., New
York, NY.

18. Zornoza, P., J. Caselles, and O. Carpena. 1987. Response of pepper plants
to NO3-:NH4+ ratio and light intensity. J. Plant Nutr. 10(7):773-782.
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