DIVISION S-4SOIL FERTILITY & PLANT NUTRITION

Salinity and Nitrogen Rate Effects on the Growth and Yield of Chile Pepper Plants
n-Valencia, and Marta D. Remmenga Magdalena Villa-Castorena, April L. Ulery,* Ernesto A. Catala ABSTRACT
Salinity and low soil N availability are important growth limiting factors for most plants. Our objective was to determine the influence of different N fertilization rates and soil salinity levels on the growth and yield of chile pepper plants (Capsicum annuum L.) grown in a greenhouse in sandy loam soil for 2 yr. The targeted soil salinity levels were 1.3, 3.5, and 5.5 dS m1 in 1999, and 1.3, 3.0, 4.5, and 6.0 dS m1 in 2000 as electrical conductivity of the saturated paste extract (ECe). Total N application rates were 80, 140, and 200 kg ha1 in 1999, and 30, 90, and 150 kg ha1 in 2000. Nitrogen rates of 140 kg ha1 or more increased soil salinities, in some cases by as much as 4 dS m1. Soil salinity decreased plant relative growth rate (RGR) up to first mature pod stage. After this growth stage however, salinity increased the RGR. Low and medium N rates produced the maximum RGR up to the first mature pod stage. After this growth stage, the maximum RGR was achieved with the medium and the high N rates tested in 1999 and 2000, respectively. Increasing N rates and salinity levels interacted to reduce chile pod yield in 1999, and acted independently in 2000. This study indicates that over-fertilization during early plant development may contribute to salinity and decreased pod yield. While salt-stressed chile performs well when adequately fertilized, N should be applied in amounts that increase with plant need over the growing season.

oil salinity is an important growth-limiting factor for most non-halophytic plants. Salts inhibit plant growth by osmotic stress, nutritional imbalance, and specific ion toxicity (Alam, 1994; Jacoby, 1994; Gunes et al., 1996; Cornillon and Palloix, 1997). Soil salinity is being progressively exacerbated by agronomic practices such as irrigation and fertilization, especially in arid regions. The proper use of N fertilizer in all soils is important, but particularly so in saline soils where N might reduce the adverse effects of salinity on plant growth and yield (Shen et al., 1994; Soliman et al., 1994; Albassam, 2001; Flores et al., 2001) depending on plant species, salinity level, or environmental conditions (Grattan and Grieve, 1999). On the other hand, over fertilization with N may contribute to soil salinization

and increase the negative effects of soil salinity on plant performance. Studies of plant growth responses to N and soil salinity during the whole plant life cycle are important to reveal whether the amount of N applied alleviates or aggravates the detrimental effects of salinity during specific growth stages. In addition, examining plant growth during the whole growing season provides information about crop salt tolerance over time. Plant response to salinity changes with age, plant development, and growth stage (Maas, 1993). For example, Arshi et al. (2002) reported that senna (Cassia angustifolia Vahl.) was most sensitive to salinity at the post-flowering stage. Relative growth rate allows one to make more appropriate comparisons of plant growth among salinity treatments than absolute growth rate (Cramer et al., 1994). The RGR gives a relative basis on which to compare growth rates of plants since it takes into account both the initial and ending plant weights over a specified time period (Hunt, 1990). The RGR is a function of the net assimilation rate (NAR), which is an index of the photosynthetic-assimilatory capacity of the plant per unit leaf area, and the leaf area ratio (LAR), which is an index of the leafiness of the plant (Hunt, 1990). Chile pepper is one of the three most important solanaceous vegetable crops in the world (Hedge, 1997) as well as being a major crop in New Mexico and other southwestern states (Johnson, 1999). It is classified as moderately sensitive to soil salinity (Maas and Hoffman, 1977). Studies on chile pepper responses to salinity and to fertilizer application under saline conditions throughout the growing season are scarce. Therefore, the objective of this work was to determine the influence of different N fertilization rates and soil salinity levels on the growth, evaluated at three different plant growth periods, and yield of chile pepper plants. MATERIALS AND METHODS
This study consisted of two greenhouse experiments conducted at New Mexico State University during the 1999 and 2000 growing seasons. Sandia chile pepper seedlings that were 6 wk old and 15 cm tall were transplanted on 29 May 1999 and 12 May 2000 into 15-L (4-gal) black polyethylene pots. This chile cultivar is long-fruited, of medium maturation time, and commonly used in local industry. Each pot was filled with 20 kg of non-saline Brazito sandy loam soil (Mixed
Abbreviations: DAT, days after transplanting; ECe, electrical conductivity of the saturated paste extract; LAR, leaf area ratio; NAR, net assimilation rate; RGR, relative growth rate; TDM, total dry matter yield.

M. Villa-Castorena and E.A. Catalan-Valencia, CENID-RASPA-INIFAP Apartado Postal No. 41, Cd. Lerdo, Dgo., Mexico 35150; A.L. Ulery, Dep. of Agronomy and Horticulture, New Mexico State University, MSC 3Q, P.O. Box 30003, Las Cruces, NM 88001; M.D. Remmenga, New Mexico State University Statistics Center, MSC 3CQ, P.O. Box 30003, Las Cruces, NM 88001. Contribution of the New Mexico Agricultural Experiment Station with additional funding by the New Mexico Chile Commission. Received 9 Nov. 2001. *Corresponding author (aulery@nmsu.edu). Published in Soil Sci. Soc. Am. J. 67:17811789 (2003). Soil Science Society of America 677 S. Segoe Rd., Madison, WI 53711 USA

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thermic Typic Torripsamment) collected from the upper 15 cm of a local agricultural field before starting each experiment in 1999 and 2000. One chile plant was grown in each pot. Different rates of N and salinity levels were tested in both years. In 1999, the rates of N were 80, 140, and 200 kg ha1, which correspond to 2.1, 3.7, and 5.4 g N per plant in terms of plant population density. These amounts of N were estimated with a population density of 37 000 plants ha1. Commercial grade NH4NO3 was used as the N source and was applied in the irrigation water in four equal quantities at 0, 15, 35, and 65 d after transplanting (DAT), according to common field practices used by local farmers. The salinity levels, measured as the ECe, were created with NaCl and CaCl2, which were dissolved in deionized water at an equivalent ratio of 1:1, and applied to each pot 1 d before transplanting. In 1999, the intended salinity levels were 1.25 (no salts added), 3.5, and 5.5 dS m1. These values changed throughout the plant-growing season due to the addition of N fertilizer and the uptake of salts by plants. There was no drainage or leaching through the pots. In the 1999 experiment, the high N rate contributed to soil salinity and had a detrimental effect on plant performance while the low and medium N rates produced similar plant responses to each other. Therefore, to investigate a lower N range, the N treatments were reduced to 30, 90, and 150 kg ha1 in the 2000 experiment, corresponding to 0.8, 2.4, and 4.0 g N plant1. An additional soil salinity level was also added for a more detailed coverage of this factor. The target ECe values tested in 2000 were 1.25, 3.0, 4.5, and 6.0 dS m1. The amount of each salt required to generate each salinity level was estimated by the following equation:

Plants were cut at the soil surface, stems and leaves were individually collected, and roots were separated from the soil with a gentle spray of tap water. Leaf area was measured for each plant with a LI-COR LI-2000 portable leaf area meter (LI-COR Inc., Lincoln, NE). Each plant component was washed with distilled water, dried in a forced air oven at 80C for 48 h, and weighed. Plant growth was evaluated in terms of RGR, and its components of NAR and LAR. These were estimated as mean values over the time interval T1 to T2 according to Hunt (1990) using the following equations:

RGR NAR

ln(W2) ln(W1) T2 T 1

[2]

(W2 W1)[(ln(LA2) ln(LA1)] (T2 T1)(LA2 LA1)

[3]

LAR 0.5

LA W

LA2 W2

[4]

(10ECe)(EW)(PSV) 2

[1]

where A was the amount of each salt (NaCl or CaCl2) added to the pot in mg, ECe was the intended or target salinity in dS m1, the constant 10 is an empirical factor used to convert ECe in dS m1 to total dissolved salts in the soil saturated paste extract in meq L1 (Dudley, 1994), EW is the equivalent weight of each salt in mg meq1, and PSV is the pot soil saturation volume in liters. Since two salts were used, we divided the numerator by 2 to consider the contribution of each salt to ECe. The PSV was calculated from soil porosity, which was estimated from the measured soil bulk density (Jury et al., 1991). A randomized complete block design with nine replications was used as the experimental design in both years. Each block included nine treatments in 1999 (81 pots), and 12 treatments in 2000 (108 pots), as a 3 3 and 3 4 factorial arrangement, respectively. At each of three growth stages, three blocks (for a total of 27 pots in 1999 and 36 pots in 2000 at each stage) were randomly selected to obtain ECe and plant data (described below). The plants were harvested at the onset of flowering, first mature pods, and onset of leaf senescence. Plant flowering onset occurred at 35 and 40 DAT; first mature pods started at 65 and 70 DAT; and leaf senescence began at 110 and 120 DAT, respectively for 1999 and 2000. Ten seedlings were collected at transplanting time to establish a control baseline for plant data. A composite soil sample was taken from each of the above selected pots to determine ECe. Five soil subsamples were collected from the whole soil pot depth with an auger, combined, and air-dried. The ECe was determined with an Accumet conductivity meter 50 (Fisher Scientific, Pittsburg, PA), after extracting solution from a saturated soil paste using the method of the U.S. Salinity Lab Staff (1954).

where LA, T, and W represent leaf area per plant (cm2), time (days), and total plant dry weight (g) respectively. Subscripts 1 and 2 refer to the values of the variable measured at two successive harvests (1 initial, 2 final). Relative growth rate is expressed in mg g1 d1, NAR in mg cm2 d1, and LAR in cm2 g1. Chile pods were collected between the second and the third growth stages to evaluate marketable yield. Plants were irrigated by hand with deionized water (EC 0.015 dS m1) every other day at the beginning of the growing season, and daily during the period of high evaporative demand. Two or three pots of each treatment were weighed daily to determine the amount of water needed to raise the soil water content to field capacity and to prevent drainage. Plant growth and yield data collected at each growth stage were subjected to analysis of variance as a factorial experiment. Because of the large number of pair-wise tests that were performed, Duncans tests at 0.05 were made to determine differences among treatments of plant dry matter accumulation, RGR, NAR, and LAR. Analysis of correlation between RGR and the variables NAR and LAR were made to determine the most influential physiological processes determining plant growth. Statistical analysis was performed using PROC GLM and PROC CORR from SAS software version 8.0 (SAS Institute Inc., 1992).

RESULTS AND DISCUSSION Nitrogen Application Rate Affects Soil Salinity

Nitrogen fertilizer treatments affected soil salinity measured as ECe (Fig. 1). Increasing N rates increased soil salinity levels measured at three plant growth stages in both years. In 1999, N rates of 140 and 200 kg ha1 increased the initial soil salinity by an average value of 0.8 and 1.5 dS m1 respectively at flowering onset, and 2 and 4 dS m1 at first mature pod stage. However, by the time of leaf senescence, soil salinity had returned to the initial values in the medium N rate while it was still 2 dS m1 above the initial values in the highest N treatment (Fig. 1). In 2000, the highest N rate of 150 kg ha1 increased soil salinity by 0.8 dS m1 at flowering onset and 1 dS m1 at first mature pod stage, but had no effect during leaf senescence (Fig. 1). This result was

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Fig. 1. Soil salinity measured as electrical conductivity of saturated soil paste (ECe) in response to targeted salinity treatments and three N application rates at (a) flowering onset, (b) first mature pod stage, and (c) leaf senescence stage in 1999 and 2000. Every point is the average of three replications and the bars are SE.

comparable with that produced by the medium N rate of 140 kg ha1 in 1999. Within the same-targeted salinity treatment, there was an increase in salinity as a result of N fertilization at rates of 140 kg ha1 and higher. The N rates of 80 and 90 kg ha1 tested in 1999 and 2000, respectively, provided chile plants with adequate N up to the first mature pod stage. Higher N rates left residual N that accumulated with that applied in subsequent fertilizations and increased temporarily the soil salinity above the targeted values. By the end of the season, N rates in excess of 150 kg ha1 contributed to soil salinities above the targeted levels. Drainage from the pots was prevented to retain the initial salts applied to the pots, which also allowed us to investigate the contribution of high fertilization rate to salinity and resultant stress. Differences were noted between 1999 and 2000 with respect to the influence of N rate on soil salinity (Fig. 1). In the 2000 experiment, the targeted and measured salinities were more in agree-

ment than in 1999 as indicated by the distance from the 1:1 line in Fig. 1. The higher average maximum temperatures in the greenhouse (35C in 1999 compared with 31C in 2000) may have exacerbated the combined osmotic effects of added salts and NH4NO3 fertilizer. To account for this microclimatic effect, results from the 2 yr will generally be reported separately in the following discussion. Excessive applications of chemical fertilizers have been reported as a factor contributing to soil salinity (Rhoades, 1990; Alam, 1994). Optimal rates of N for maximum chile pepper yields vary from 70 (OSullivan, 1979) to 180 kg ha1 (Moreno et al., 1996), depending on soil N availability. We tested N rates under and over these recommended levels and found that 200 kg ha1 resulted in salt accumulation at the end of the growing season. Thus, fertilization above plant requirements not only increases costs, but can also have detrimental effects on the environment such as soil salinity accumula-

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Table 1. Total plant dry matter (TDM in g per plant) as affected by soil salinity and N rate at three plant growth periods in 1999 and 2000.
Year Plant growth period Target soil salinity dS 1999 At flowering onset 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 6.8 4.7 4.4 5.3 a 75.5 44.5 25.3 48.4 a 105.8 95.4 74.7 92.0 a 30 kg ha1 At flowering onset 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 13.7 12.9 9.4 8.1 11.0 b 24.2 23.0 20.5 16.6 21.1 c 55.4 49.8 45.4 44.4 48.8 c 9.2 2.6 1.9 4.6 ab 56.1 16.8 8.9 27.3 b 117.4 103.4 47.9 89.6 a 90 kg ha1 16.7 14.4 12.8 10.9 13.7 a 49.5 31.8 35.3 31.7 37.1 a 91.5 111.3 95.5 95.5 98.5 b 5.6 2.5 1.7 3.3 b 39.6 6.0 5.5 17.0 c 100.0 60.6 30.3 63.6 b 150 kg ha1 10.6 9.0 8.3 7.0 8.7 c 38.6 31.0 24.5 22.7 29.2 b 153.6 137.4 109.4 91.1 122.9 a 13.7 12.1 10.1 8.7 37.5 28.6 26.8 23.7 100.2 99.5 83.5 77.0 a ab bc c a b b b a a b b 7.2 a 3.3 b 2.7 b 57.1 a 22.4 b 13.2 c 107.7 a 86.5 b 51.0 c m 1 Total N applied 80 kg ha1 140 kg ha1 g plant1 200 kg ha1 Mean

At first mature pods

At leaf senescence

2000

At first mature pods

At leaf senescence

Means followed by the same letter are not significantly different within rows and column according to Duncans test (P 0.05, n 3) for each year and growth stage.

tion or conversely, ground water contamination by leaching.

Soil Salinity and Nitrogen Effects on Total Plant Dry Matter Accumulation
Salinity and N rate both affected total plant dry matter (TDM) accumulation at flowering onset, first mature pods, and leaf senescence growth stages in 1999. However, no interaction between salinity and N rate was detected, that is, the effects caused by the N rates did not depend on the salinity levels and vice versa. Salinity reduced TDM at all three growth stages (Table 1), but at the end of the growing season, at leaf senescence, the relative TDM reductions caused by salinity were lower than those produced at earlier growth stages. Thus it appears that the observed recovery of plant growth toward the end of the plant cycle is a possible mechanism of plant adaptation to soil salinity. Increasing amounts of N had a detrimental effect on TDM in all plant growth stages in 1999 due to the contribution of N fertilizer to soil salinity as described in the previous section (Fig. 1). This effect was more pronounced at the first mature pods stage when the contribution of fertilizer to soil salinity, specifically N rates of 140 and 200 kg ha1, was highest. Thus, high N rates applied during early plant development may have detrimental effects on dry matter production of chile grown in saline conditions. Salinity and N rate effects on TDM were also observed for the 2000 growing season. No interaction between these two factors was detected at the flowering

onset or first mature pods stage, but at leaf senescence, an interaction was found. The TDM was reduced by salinity levels of 4.5 dS m1 and above at flowering onset and leaf senescence and by salinity levels higher than or equal to 3.0 dS m1 at first mature pods stage (Table 1). However, the TDM decrease due to soil salinity was less in 2000 than in 1999 and may have been because of lower air temperatures in the greenhouse. The effects of N rate on TDM differed with plant growth stage. The N rate of 90 kg N ha1 produced the maximum TDM value at flowering onset and first mature pod stages. On the other hand, TDM increased with increasing N rates during leaf senescence, except for the highest salinity level of 6.0 dS m1.

Soil Salinity and Nitrogen Effects on Green Pod Yield

In 1999, both salinity and N rate affected pod yield and an interaction was detected between these factors (Fig. 2). Target salinities of 1.25 and 3.5 dS m1 had similar pod yields, which were higher than the yield obtained with 5.5 dS m1 at the lowest N rate of 80 kg ha1. However, increasing salinity reduced pod yield much more in soils treated with higher rates of N such as 140 and 200 kg ha1. This was because high N rates increased soil salinity beyond the targeted values by up to 2 and 4 dS m1, respectively, as previously discussed. In 2000, when the applied N rates were lower than those tested in 1999, salinity and N rate also affected pod yields but an interaction between them was not

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Fig. 2. Chile green pod yield response to soil salinity as affected by N rate applied in 1999 and 2000. Every point is the mean from three replications and the bars are SE. The arrows indicate the target salinity levels for each experiment.

detected. Target salinities of up to 4.5 dS m1 caused no decrease in pod yield, but at 6 dS m1 yields were significantly lower. Contrary to 1999, increasing N rates resulted in increased pod yields in the 2000 experiment. Soil salinity decreased chile pod yield more in 1999 than in 2000 at similar N rates (Fig. 2). That is, the decrease in pod yield as a function of ECe was more abrupt in 1999 for plants treated with 80 or 140 kg N ha1 than in 2000 when comparable N rates of 90 or 150 kg ha1 were applied. The difference in greenhouse temperature for the two growing seasons was the only parameter observed that might have caused this exaggerated osmotic effect as it is known that plant response to salinity varies with many factors such as climate, soil conditions, agronomic practices, and stage of growth (Rhoades, 1990; Maas, 1993).

tion between these two factors detected (Table 2). During the first growth period, from transplanting to flowering onset, salinity levels greater than or equal to 3.5 dS m1 decreased the RGR. From flowering onset to first mature pods, the RGR was reduced by the target salinity ECe of 5.5 dS m1. Conversely, salinity had a positive effect on RGR in the last development period, from first mature pods to leaf senescence. However, this apparent beneficial effect of salinity on RGR late in the plants life cycle may be because salt-stressed plants delayed their early growth and development and subsequently continued growing longer into the season, resulting in a higher RGR while nonstressed plants had already reached leaf senescence. It is known that salinity stress retards plant growth due to osmotic and specific ion effects on metabolic processes such as nutrient uptake and photosynthesis (Alam, 1994). Increasing N rates from 80 to 200 kg ha1 also reduced the RGR during the early and middle development periods. In contrast, N rates of 140 and 200 kg ha1 both increased the RGR with respect to 80 kg ha1 toward the end of the growing cycle, from first mature pods to leaf senescence period (Table 2). In 2000, soil salinity reduced the RGR during early plant development but then caused an increase in RGR during the last development period (Table 2). Soil salinity of 4.5 dS m1 or above decreased the RGR during transplanting to flowering onset, but salinities of 3.0 dS m1 or more tended to increase the RGR from first mature pods to leaf senescence. These results are similar to those found in 1999 with respect to differences in salt tolerance among plant growth periods. However, the salinity levels that were required to reduce the RGR in 2000 were higher than those in 1999 possibly due to temperature differences as discussed earlier. Other studies have also pointed out the strong influences of air temperature on the responses of plants to soil salinity (Maas, 1993). There was no interaction detected between soil salinity and N rate for any of the plant growth periods in 2000 (Table 2). Increasing the N rate from 30 to 90 kg ha1 increased the RGR during transplanting to flowering onset and from flowering onset to first mature pods. But at the end of growing season, an increase from 30 to 150 kg N ha1 was needed to produce the highest RGR. These results were in agreement with those found in 1999 when 80 and 140 kg N ha1 were enough to satisfy plant N requirements from transplanting to first mature pod and from first mature pod to leaf senescence periods, respectively. Thus, it may be possible to improve chile plant yields by supplying N fertilizer in amounts that increase with plant size instead of in four equal application rates since salt stress was intensified by over-fertilization during times of low fertilizer requirements.

Soil Salinity and Nitrogen Effects on Relative Growth Rate

The RGR was affected by soil salinity and N rate during all plant growth periods in 1999 with no interac-

Soil Salinity and Nitrogen Effects on Net Assimilation Rate

Soil salinity affected the NAR throughout the growing season in 1999 but N rate only affected NAR after

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Table 2. Relative growth rate (RGR in mg g1 d1) as affected by soil salinity and N rate over three plant growth periods in 1999 and 2000.
Total N applied Year 1999 From transplanting to flowering onset 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 105.4 88.0 93.2 95.5 a 82.1 84.5 58.2 74.9 a 6.2 14.0 20.7 13.6 b 30 kg ha1 From transplanting to flowering onset 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 89.6 88.2 80.8 77.8 84.1 b 22.0 23.22 30.71 28.27 26.06 b 20.89 19.29 20.52 24.77 21.37 b 115.5 76.0 70.5 87.4 ab 60.4 65.6 46.8 57.6 ab 13.3 33.1 32.9 26.5 a 90 kg ha1 93.7 90.6 88.0 84.3 89.1 a 43.9 31.61 40.61 42.55 39.66 a 15.40 31.28 24.96 27.88 24.88 b 99.4 77.6 67.8 81.6 b 67.6 28.7 33.9 43.4 b 16.4 42.7 33.1 30.8 a 150 kg ha1 82.9 79.2 78.0 74.6 78.7 c 53.5 50.49 44.05 46.07 48.52 a 34.45 37.77 37.45 35.44 36.28 a 88.7 86.0 82.2 78.9 a ab bc c 106.8 a 80.5 b 77.2 b 70.0 a 59.6 ab 46.3 b 12.0 b 30.0 a 30.8 a Plant growth period Target soil salinity dS m 1 80 kg ha1 140 kg ha1 mg g1 d 1 200 kg ha1 Mean

From flowering onset to first mature pods

From first mature pods to leaf senescence

2000

From flowering onset to first mature pods

39.8 a 35.11 a 38.46 a 38.96 a 23.58 29.45 27.64 29.37 b a ab a

From first mature pods to leaf senescence

Means followed by the same letter are not significantly different within rows and column according to Duncans test (P 0.05, n 3) for each year and development period.

the onset of flowering (Table 3). No interaction between salinity and N rate was detected for any plant growth period. Target salinities of 3.5 and 5.5 dS m1 reduced NAR by about 30% compared with the unsalinized treatment during the period of transplanting to flowering onset. These salinity levels also reduced NAR by approximately 35% compared with unsalinized treatments from flowering onset to first mature pods stage. In contrast, perhaps as a consequence of plant growth delay caused by salinity, the NAR was increased about two-fold by the salinity treatment of 3.5 and 5.5 dS m1 during the last development period. The NAR, which represents a balance between the physiological processes of photosynthesis and respiration of the whole plant, decreases due to reductions in photosynthesis or increases in maintenance respiration (Cramer et al., 1990). Several studies have shown reductions in photosynthesis due to salt stress (Myers et al., 1990; Tiwari et al., 1997), which has been attributed to decreased stomatal, and mesophyll conductance to CO2 diffusion (Delfine et al., 1999) as well as the inhibition of biochemical reactions and reduced carboxylase activity (Ungar, 1991). In addition, reduced photosynthesis in saline conditions has been correlated with leaf Na or Cl accumulation (Bethke and Drew, 1992). There is also evidence that salt stress increases maintenance res lez-Moreno et al., 1997) which is mainly piration (Gonza attributed to additional energy expenditures required for the salt economy of the cell, that is, pumping out ions from the cytosol into the vacuole (Semikhatova et al., 1993).

In 1999, increasing N rates from 80 to 200 kg ha1 reduced NAR during flowering onset to the first mature pod stage (Table 3). This indicates that over-fertilization is detrimental for plant growth and may be due to the osmotic effect caused by added fertilizer. In contrast, during the period of first mature pods to leaf senescence, the NAR positively responded to increases in N rate from 80 to 140 or 200 kg ha1. In 2000, the NAR was not affected by salinity at any plant growth period. The NAR did increase in response to N rates of 150 kg ha1 during the two latter growth stages, flowering onset to leaf senescence.

Soil Salinity and Nitrogen Effects on Leaf Area Ratio

Salinity and N rate affected LAR primarily during the first mature pod to leaf senescence period in 1999 (Table 4). No interaction was detected between salinity and N rate. The targeted salinity treatment of 5.5 dS m1 increased the LAR by 34% with respect to the unsalinized treatment. Salinity has also been shown to induce higher LAR in sweetclover (Melilotus segetalis n, 1994). In addition, Brot Ser.) (Romero and Maran o an increase of similar magnitude in LAR was also produced by the high N rate of 200 kg ha1. In 2000, there was no effect due to salinity during the first plant growth stage (Table 4). From flowering onset to first mature pods, only the 3.0 dS m1 salinity treatment increased LAR with respect to the less saline treatment. During the last growth stage, salinity treatments

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Table 3. Net assimilation rate (NAR in mg cm2 d1) as affected by soil salinity and N rate over three development periods in 1999 and 2000.
Total N applied Year 1999 From transplanting to flowering onset 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.08 0.98 0.95 1.00 a 1.53 1.15 1.04 1.24 a 0.23 0.64 0.69 0.52 b 30 kg ha1 From transplanting to flowering onset 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 1.00 1.19 1.16 1.02 1.09 a 0.32 0.36 0.49 0.43 0.40 b 0.56 0.47 0.48 0.57 0.52 b 1.53 0.68 0.76 0.99 a 1.10 0.96 0.68 0.91 ab 0.55 1.25 0.86 0.89 a 90 kg ha1 1.18 1.07 1.04 1.12 1.10 a 0.60 0.37 0.49 0.56 0.51 ab 0.38 0.71 0.59 0.62 0.58 b 1.19 0.96 0.87 1.00 a 1.11 0.49 0.58 0.73 b 0.50 1.07 0.83 0.80 a 150 kg ha1 1.11 0.86 0.99 1.04 1.00 a 0.68 0.55 0.52 0.57 0.58 a 0.85 0.87 0.82 0.74 0.82 a 1.09 1.04 1.06 1.06 0.53 0.43 0.50 0.52 0.59 0.69 0.63 0.65 a a a a a a a a a a a a 1.27 a 0.87 b 0.86 b 1.25 a 0.87 b 0.77 b 0.42 b 0.99 a 0.80 a Plant growth period Target soil salinity dS m1 80 kg ha1 140 kg ha1 mg cm2 d1 200 kg ha1 Mean

From flowering onset to first mature pods

From first mature pods to leaf senescence

2000

From flowering onset to first mature pods

From first mature pods to leaf senescence

Means followed by the same letter are not significantly different within rows and column according to Duncans test (P 0.05, n 3) for each year and development period.

Table 4. Leaf area ratio (LAR in cm2 g1) as affected by soil salinity and N rate over three development periods in 1999 and 2000.
Total N applied Year 1999 From transplanting to flowering onset 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.3 3.5 5.5 Mean 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 1.3 3.0 4.5 6.0 Mean 112.38 116.73 94.77 107.96 a 66.44 78.05 57.27 67.25 a 28.75 32.66 37.49 32.97 b 30 kg ha1 97.84 88.43 85.29 89.56 90.28 a 71.44 65.18 63.93 67.65 67.05 b 42.23 45.99 48.09 49.03 46.34 c 98.96 125.21 106.70 110.29 a 58.29 72.10 75.89 68.76 a 31.25 32.25 48.99 37.50 b 90 kg ha1 92.57 95.55 94.94 89.25 93.08 a 73.31 84.66 82.28 75.57 78.96 a 48.23 57.42 54.54 55.34 53.88 b 102.78 99.39 96.55 99.57 a 63.94 63.63 61.34 62.97 a 38.45 46.75 45.80 43.67 a 150 kg ha1 89.25 99.79 90.51 86.67 91.056 a 78.30 90.63 82.63 79.65 82.80 a 56.02 58.77 61.17 62.60 59.64 a 104.71 ab 113.77 a 99.34 b 62.89 a 71.26 a 64.83 a 32.82 b 37.22 b 44.09 a Plant growth period Target soil salinity dS m1 80 kg ha1 140 kg ha1 cm2 g1 200 kg ha1 Mean

From flowering onset to first mature pods

From first mature pods to leaf senescence

2000 From transplanting to flowering onset

93.22 94.59 90.25 88.49 74.35 80.15 76.28 74.29 48.83 54.06 54.60 55.66

ab a ab b b a ab b b a a a

From flowering onset to first mature pods

From first mature pods to leaf senescence

Means followed by the same letter are not significantly different within rows and columns according to Duncans test (P 0.05, n 3) for each year and development period.

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Table 5. Correlation coefficients between relative growth rate (RGR) and net assimilation rate (NAR), and between (RGR) and leaf area ratio (LAR) at three growth periods for 2 yr. Each correlation value was obtained from 9 and 12 pairs of data (number of salinity levels times three replications for 1999 and 2000, respectively).
Year 1999 Development period Transplanting to first flowers Flowering onset to first mature pods First mature pods to leaf senescence 2000 Transplanting to first flowers Flowering onset to first mature pods First mature pods to leaf senescence N rate applied 80 140 200 80 140 200 80 140 200 30 90 150 30 90 150 30 90 150 NAR 0.65 0.91 0.83 0.49 0.91 0.98 0.81 0.89 0.98 0.16 0.60 0.50 0.99 0.96 0.95 0.91 0.98 0.74 NS ** ** NS ** ** ** ** ** NS * NS ** ** ** ** ** ** LAR 0.59 0.61 0.21 0.79 0.21 0.13 0.88 0.57 0.85 0.51 0.13 0.01 0.63 0.39 0.03 0.67 0.79 0.62 NS NS NS ** NS NS ** NS ** NS NS NS ** NS NS ** ** *

NS not significant or only significant at P 0.05. * Significant at 0.01 P 0.05 ** Significant at P 0.01.

of 3.0 dS m1 or more caused an increase in LAR. The N rates of 90 and 150 kg ha1 increased the LAR during the two last development periods compared with 30 kg N ha1.

Correlations between Growth Parameters

The NAR and RGR were highly correlated for most of the N rates and plant growth periods (Table 5). Thus, changes in RGR can be explained by concomitant decreases or increases in photosynthesis per leaf area and respiration reflected in the NAR. Salt stressed barley plants (Hordeum vulgare) (Cramer et al., 1990) and n, 1994) were also sweet clover (Romero and Maran o shown to have high correlation between NAR and RGR. The correlation between LAR and RGR was considerably lower than that between NAR and RGR for most of the N rates and plant growth periods (Table 5). An exception was the greater correlation between LAR and RGR at 80 kg N ha1 during reproductive growth in 1999. This suggests that leaf expansion was not limiting the RGR of most of the plants affected by salt stress. However, leaf expansion appeared to contribute to high RGR values in salt-stressed plants during the period of first mature pods to leaf senescence. The LAR has been shown to be highly correlated with the RGR of saltstressed maize (Zea mays) (Cramer et al., 1994) and n, 1994). But LAR sweet clover (Romero and Maran o was not correlated with the RGR of salt-stressed barley plants (Cramer et al., 1990). Munns and Termaat (1986) proposed that salt stress acts by increased leaf death rates, not decreased NAR. The ability to maintain functional leaf area by increasing LAR, as seen here, can be adaptive.

plants were apparently aggravated by higher average maximum temperatures in the greenhouse in 1999 (35C) compared with 2000 (31C). Thus increasing N rates and salinity levels interacted to reduce chile pod yield in 1999, but acted independently in 2000 when lower N rates were applied and greenhouse temperatures were lower. The RGR decreased in response to salinity early in the growing season, but was higher during later growth stages because salt-stressed plants delayed their early growth and development and subsequently continued growing longer into the season while non salt-stressed plants had already reached leaf senescence. However the recovery in RGR later in the plants life cycle did not compensate for the final pod yield. The N rates of 80 and 90 kg ha1 tested in 1999 and 2000 provided the plant with enough N to produce the maximum RGR up to the first mature pod stage while higher N rates tended to reduce RGR. After the first mature pods stage, the maximum RGR was achieved with the N rates of 140 and 150 kg ha1 in 1999 and 2000, respectively. Salt-stressed chile pepper plants can grow and produce high pod yields when adequately fertilized by supplying N in amounts that increase with plant size. The addition of excess N as NH4NO3 fertilizer did not alleviate osmotic stress as hypothesized by some authors, and in fact, contributed to the overall soil salinity as measured by ECe. Therefore, over-fertilization early in the plant life cycle can be both destructive and wasteful. REFERENCES
Alam, S.M. 1994. Nutrient uptake by plants under stress condition. p. 227243. In M. Pessarakli (ed.) Handbook of plant and crop stress. Marcel Dekker, New York. Albassam, B.A. 2001. Effect of nitrate nutrition on growth and nitrogen assimilation of pearl millet exposed to sodium chloride stress. J. Plant Nutr. 24:13251335. Arshi, A., A.Z. Abdin, and M. Iqbal. 2002. Growth and metabolism of senna as affected by salt stress. Biol. Plant. 45:295298. Bethke, P.C., and M.C. Drew. 1992. Stomatal and nonstomatal components to inhibition of photosynthesis in leaves of Capsicum annuum

CONCLUSIONS
Soil salinity decreased plant growth and yield of Sandia chile pepper plants grown for two separate years in a greenhouse. High N fertilization rates contributed to soil salinity. The negative effects of salinity on the

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