The objectives of these greenhouse experiments were to determine the effects of elevated CO2 on growth, mineral nutrition, and gas exchange physiology of seedlings of four commercial Citrus rootstocks. We grew well-watered and fertilized seedlings of `Volkamer' lemon (VL), `Cleopatra' mandarin (CL), `Swingle' citrumelo (SW), and `Troyer' citrange (TC) cultivars (in decreasing order of vigor) in unshaded, air-conditioned greenhouses at ambient CO2 (350 μmol/mol) or 2x ambient CO2 for 5 months. CL was the smallest cultivar, had the lowest root/shoot (r/s) ratio,and lowest rates of CO2 assimilation (A) of leaves, transpiration (E), and water-use efficiency, (A/E). Overall, daily whole-plant water use was correlated with single-leaf E. Elevated CO2 increased both shoot and root growth similarly; therefore, r/s was not affected. Elevated CO2 increased A, leaf dry wt/area, and leaf C, but decreased transpiration and leaf N so that leaf C/N, A/N, and A/E all increased. Although plant size of the four cultivars ranked similarly at both ambient and high CO2, the more-vigorous cultivars grew proportionately more at high CO2 than the less-vigorous cultivars. Growing cultivars at elevated CO2 can yield insights into mechanisms determining vigor and relationships between A and plant growth.
J.P. Syvertsen and M. Salyani
The effects of three highly refined petroleum spray oils and of ambient vapor pressure on net CO2 assimilation (A) and stomatal conductance of water vapor (gs) of single grapefruit (Citrus paradisi Macf.) leaves were investigated. Overall, gs of various-aged leaves was decreased by a large leaf-to-air vapor pressure difference (VPD). In the first experiment, oils with midpoint distillation temperatures (50% DT) of 224, 235, and 247C were applied with a hand atomizer at concentrations of 0, 1%, and 4% oil emulsions in water and 100% oil, all with 0.82% surfactant (by volume). There was a tendency for oils of the two higher DT to decrease net gas exchange during a subsequent 12 days, but significant differences could not be attributed to oil DT. Both A and gs were reduced by the two higher concentrations of oil mixtures. In the second experiment, a commercial airblast sprayer was used to apply the 224C oil at 4% or the 235C oil at 2% and 4% mixtures plus surfactant under field conditions. There were no significant effects of oil treatments on net gas exchange of leaves either measured under moderate VPD outdoors 1 day after spraying or under low VPD in the laboratory 2 days after spraying. No visible phytotoxic symptoms were observed in either experiment.
J.P. Syvertsen and J.M. Dunlop
We tested the hypothesis that amendments of two hydrophilic gels to a sand soil would reduce N leaching losses and increase growth of citrus seedlings. Three-month-old seedlings of `Swingle' citrumelo [Citrus paradisi Macf. × Poncirus trifoliata (L.) Raf.] were transplanted into containers of steam-sterilized Candler sand, amended with a linear acrylamide/acrylate copolymer (PAM), and/or a cross-linked copolymer agronomic gel (AGRO). Two rates of each amendment were applied either alone or together and were either mixed into dry sand prior to seedling transplant, used as a root-dip slurry at transplant or applied to the soil surface in a solution after transplant. Seedlings were grown in the greenhouse for 5 months and irrigated to container capacity with a dilute nutrient solution without leaching. Pots were leached every 2 weeks and total N losses from the soil were measured in the leachate. PAM amendments increased N retention in soil slightly but PAM had no effect on plant growth, water use, N uptake, or N leaching relative to unamended control plants. The AGRO amendments increased seedling growth, plant water use and uptake of N from 11% to 45% above that of the unamended control plants depending on application method. AGRO decreased N concentrations in the leachate to as low as 1 to 6 mg·L-1. Only 6% of the total applied N was leached from the AGRO treatments, which was about half that from the untreated control plants. There was no additional benefit of using both amendments together or of an additional AGRO root dip treatment. The largest plants used the most water, required the most N and had the greatest N uptake efficiency. AGRO amendments clearly enhanced seedling growth, increased their N uptake efficiency, and reduced N losses from this sand soil.
R. Romero-Aranda and J.P. Syvertsen
We compared net gas exchange rates of CO2 and H2O vapor of greenhouse-grown `Duncan' grapefruit (Citrus paradisi Macf.) and `Valencia' orange [C. sinensis (L.) Osbeck] leaves after multiple foliar sprays of urea N with and without NaCl: CaCl2 solutions. Highly saline solutions (3.8 dSm-1) caused necrotic burn symptoms after leaf chloride levels reached 7 mmol·m-2. Grapefruit leaves had higher leaf Cl and more burn symptoms than orange leaves. The remaining green areas of all salt-stressed leaves, however, had similar rates of net CO2 assimilation (ACO2) and stomatal conductance (gs) as water-sprayed control leaves. Total leaf N and chlorophyll increased with repeated foliar applications of urea solutions regardless of salinity levels in the spray solution. Thus, salts in solution did not interfere with foliar absorption of N. High urea N solutions (33.6 g·liter-1) without salts caused foliar burn and leaf abscission after one application. Three sprays of urea-N solution (11.2 g·liter-1) increased N concentration of N-deficient leaves about 60% and increased ACO2 rate about 50%. ACO2 did not increase when nitrogen concentration in leaves exceeded a threshold value of about 200 mmol·m-2 so photosynthetic nitrogen use efficiency (PNUE = ACO2/N) decreased with increasing leaf N concentration. Net gas exchange and PNUE was higher for grapefruit than for orange leaves. Leaf Cl levels from foliar-applied salts may not be as detrimental to leaf gas exchange as Cl from salts in soil-applied irrigation water.
J.P. Syvertsen and M.L. Smith
Four-year-old `Redblush' grapefruit (Citrus paradisi Macf.) trees on either the relatively fast-growing rootstock `Volkamer' lemon (VL) (C. volkameriana Ten. & Pasq.) or on the slower-growing rootstock sour orange (SO) (C. aurantium L.) were transplanted into 7.9-m3 drainage lysimeter tanks filled with native Candler sand, irrigated similarly, and fertilized at three N rates during 2.5 years. After 6 months, effects of N application rate and rootstock on tree growth, evapotranspiration, fruit yield, N uptake, and leaching were measured during the following 2 years. When trees were 5 years old, low, medium, and high N application rates averaged about 79,180, or 543 g N/tree per year and about 126,455, or 868 g N/tree during the following year. Recommended rates average about 558 g N/tree per year. A lysimeter tank with no tree and additional trees growing outside lysimeters received the medium N treatment. Nitrogen concentration in the drainage water increased with N rate and exceeded 10 mg·liter-1 for trees receiving the high rates and also for the no tree tank. Leachate N concentration and total N recovered was greater from trees on SO than from those on VL. Average N uptake efficiency of medium N rate trees on VL was 6870 of the applied N and 61 % for trees on SO. Nitrogen uptake efficiency decreased with increased N application rates. Trees outside lysimeters had lower leaf N and fruit yield than lysimeter trees. Overall, canopy volume and leaf N concentration increased with N rate, but there was no effect of N rate on fibrous root dry weight. Fruit yield of trees on SO was not affected by N rate but higher N resulted in greater yield for trees on VL. Rootstock had no effect on leaf N concentration, but trees on VI. developed larger canopies, had greater fibrous root dry weight, used more water, and yielded more fruit than trees on SO. Based on growth, fruit yield and N leaching losses, currently recommended N rates were appropriate for trees on the more vigorous VL rootstock but were 22% to 69 % too high for trees on SO.
Francisco García-Sánchez and J.P. Syvertsen
Three-month-old citrus rootstock seedlings of the Cl- excluder Cleopatra mandarin (Citrus reticulata Blanco) and the Cl- accumulator Carrizo citrange [C. sinensis (L.) Osb. × Poncirus trifoliata L.] were fertilized with nutrient solution with or without additional 50 mm NaCl and grown at either ambient CO2 (360 μL·L-1) or elevated CO2 (700 μL·L-1) in similar controlled environment greenhouses for 8 weeks. Elevated CO2 increased plant growth, shoot/root ratio, leaf dry weight per area, net assimilation of CO2, chlorophyll, and water-use efficiency but decreased transpiration rate. Elevated CO2 decreased leaf Ca2+ and N concentration in non-salinized Cleopatra. Salinity increased leaf Cl- and Na+ in both genotypes. Carrizo had higher concentrations of Cl-but lower Na+ in leaves than Cleopatra. Salinity decreased plant growth, shoot/root ratio, net gas exchange, water use, and root Ca+2 but increased root N in both genotypes regardless of CO2 level. Neither salinity nor elevated CO2 affected leaf chlorophyll fluorescence (Fv/Fm). Carrizo had higher Fv/Fm, leaf gas exchange, chlorophyll, N, and Ca2+ than Cleopatra. Salinity-induced decreases in leaf osmotic potential increased leaf turgor especially at elevated CO2. The increase in leaf growth at elevated CO2 was greater in salinized than in nonsalinized Carrizo but was similar in Cleopatra seedlings regardless of salt treatment. In addition, salinity decreased water-use efficiency more at elevated CO2 than at ambient CO2 in Cleopatra but not in Carrizo. Elevated CO2 also decreased leaf Cl- and Na+ in Carrizo but tended to increase both ions in Cleopatra leaves. Based on leaf growth, water-use efficiency and salt ion accumulation, elevated CO2 increased salinity tolerance in the relatively salt-sensitive Carrizo more than in the salt-tolerant Cleopatra. In salinized seedlings of both genotypes, Cl- and Na+ concentration changes in response to eCO2 in leaves vs. roots were generally in opposite directions. Thus, the modifications of citrus seedling responses to salinity by the higher growth and lower transpiration at elevated CO2 were not only species dependent, but also involved whole plant growth and allocations of Na+ and Cl-.
D. M. Eissenstat and J. P. Syvertsen
The effects of elevated levels of ozone on growth, mineral nutrition and freeze resistance were studied using broadleaf-evergreen citrus and avocado trees. `Ruby Red' grapefruit (Citrus paradisi L.) trees on either Volkamer lemon (Citrus volkameriana Ten. & Pasq.) or sour orange (Citrus aurantium L.) rootstock and `Simmonds' or `Pancho' avocado trees (Persea americana Mill.) on the rootstock `Waldin' were exposed to ozone in open-top chambers for 4 mo in 1988 and in a second experiment in 1989 for 8 mo. Citrus tree growth, estimated by total leaf mass, was unaffected by ozone concentrations of 3 times ambient in either year but avocado growth was reduced by ozone concentration at 2 times ambient in 1989. All trees were well-fertilized and ozone had little effect on mineral nutrient concentrations in leaves. Freeze resistance, estimated by electrolyte leakage from leaf disks and survival of leaves, stems, and whole-plants following exposure to freezing temperatures, was often diminished in avocado and citrus at 3 times ambient ozone, but occasionally was increased at 2 times ambient. Thus, ozone can be related to shifts in freeze resistance that can occur prior to discernible growth effects.
R. Romero-Aranda and J.P. Syvertsen
The penetration of foliar-applied urea and salt solutions into citrus leaves was investigated using `Duncan' grapefruit and `Valencia' orange seedlings in a greenhouse, and 8-year-old `Ruby Red' grapefruit trees in field tests during the summer and fall. Net gas exchange rates, Cl, nitrogen, and chlorophyll concentrations of singles leaves were measured during or after the period of foliar applications. Foliar-applied salt treatments increased leaf Cl, and visible burn symptoms were observed when Cl levels reached ≈0.4% of leaf dry weight. After 11 weeks, green areas from salt-treated leaves had similar rates of net CO2 assimilation as control plants. Leaf nitrogen and total chlorophyll increased with repeated sprays. Urea sprayed at 15% caused foliar burn symptoms after two to three applications and increased the amount of leaf abscission. Urea sprayed at 6% increased CO2 assimilation rate ≈50% after three foliar applications in 3 weeks. Nitrogen content and net CO2 assimilation of urea and urea + salt leaves were similar.
J.P. Syvertsen, M.L. Smith, and B.J. Boman
Effects of salinized irrigation water on tree canopy and root growth, water use, foliar nutrition, and leaching losses below the rootzone were studied during a 2-year period using single tree lysimeters. Eighteen 6-year-old `Valencia' orange trees on either Carrizo citrange (CC) rootstock or sour orange (SO) rootstock were each transplanted into 7.8 m3 drainage lysimeters and irrigated with water having an electrical conductivity of 0.3, 1.6, or 2.5 dS m-1 from a 3:1 ratio of NaCl:CaCl2. Six additional trees (3 on each rootstock) were transplanted into soil without tanks. Trees outside the tanks were smaller, but nutritionally similar to the low salinity trees in lysimeters. Trees on CC were larger, had greater root densities, and were associated with less leaching of ions and nutrients into drainage water from the tanks than trees on SO. High salinity irrigation water reduced canopy growth and ET, but increased fibrous root dry weight. Trees on CC accumulated more Cl in leaves and in fruit juice than those on SO. Leaching loss of total N varied from 2-8% of that annually applied to trees, but up to 70% of the applied N and up to 80% of the applied K were leached from the blank tank with no tree. Salinized trees lost more N and K to drainage water, especially those on SO. Tree size, root density, and irrigation water quality can influence leaching losses beyond the rootzone.
J.P. Syvertsen, L.S. Lee, and J.W Grosser
Diploid (2x) and autotetraploid (4x) Citrus L. rootstock cultivars were grown at elevated CO2 to obtain insights into limitations on growth and net gas exchange that have been associated with tetraploidy. Well-nourished 2x and 4x seedlings of `Volkamer' lemon (Volk, C. volkameriana Ten & Pasq.), `Troyer' citrange [Troy, C. sinensis (L.) Osbeck × Poncirus trifoliata (L.) Raf.] and `Cleopatra' mandarin (Cleo, C. reticulata Blanco.), were grown in greenhouses at either ambient or twice ambient CO2 for 4 months. Plant growth, water relations, mineral nutrition, and net gas exchange characteristics of leaves were measured. Most 4x plants were smaller and had lower rates of whole plant transpiration but shorter fibrous roots than 2x plants. Fibrous roots of 4x were thicker than 2x roots as indicated by a lower specific root length (SRL) in 4x than in 2x roots. Root hydraulic conductivity was correlated to total plant growth but there were no effects of CO2 or ploidy on root conductivity. Tetraploid leaves had lower N concentrations than 2x leaves when expressed on a dry weight basis but these differences disappeared when N concentration was expressed on an leaf area basis because 4x leaves had more leaf dry weight per area (LDW/a) than 2x leaves. Plant growth was greater and SRL was lower at elevated CO2 than at ambient CO2. LDW concentrations of N, P, and K were lower at elevated CO2 than at ambient apparently due to a growth dilution effect. LDW/a, net CO2 assimilation (ACO2), and leaf water use efficiency were greater at elevated CO2 than at ambient. Overall, there was no effect of ploidy on ACO2 but 4x Volk and Troy had lower rates of ACO2 than their 2x at elevated CO2. Net gas exchange of tetraploid leaves was less responsive to elevated CO2 than 2x leaves. The low SRL of tetraploids was correlated with low whole plant transpiration rates and low leaf area-based N concentrations, which may be operative in determining the growth characteristics associated with tetraploidy.