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- Author or Editor: Royal G. Fader x
The plant cuticle is the prime barrier to penetration of foliar-applied plant growth regulators (PGR). Spray additives of various chemistries are frequently included in a tank mix to increase performance of PGRs. We have reported that urea and ammonium nitrate (AN) enhance transcuticular penetration of 14C-labeled NAA (pKa 4.2) from aqueous droplets (pH 5.2) and their subsequent deposits through enzymatically isolated tomato fruit cuticular membranes (CM). Studies on effects of Triton × surfactants on AN-enhanced NAA penetration showed an additional 25% increase in NAA penetration and the AN:surfactant interaction was significant. Also, some alkylamine hydrochlorides increased NAA penetration. Studies comparing NAA penetration through tomato and pepper fruit and Citrus leaf CM in the presence of 8 mM AN or 8 mM ethylamine HCl showed that all three species exhibited the same trend for penetration at 120 h: ethylamine HCl > AN > NAA only. Comparative NAA penetration for CM of the three species was pepper > Citrus > tomato, with significant differences (P > 0.006) in NAA penetration, as indexed by initial slope and penetration after 120 h. On addition of AN, NAA penetration was greater (range 3% to 40%) for Citrus and pepper CM than tomato CM. When ethylamine HCl was added, NAA penetration through Citrus and pepper CM was less (–37 and –27%, respectively) than tomato CM as measured by the initial slope, but 6% and 11%, respectively, more than tomato CM for penetration after 120 h. The differences in NAA penetration among the three species cannot be explained by cuticle thickness, since pepper and tomato CM are 2.5- to 3.5-fold thicker than Citrus CM. We have suggested that the enhanced NAA penetration mediated by AN and ethylamine HCl (and other alkylamine HCl examined) may be related to their hygroscopic properties leading to greater deposit hydration. The significance of the differences among the species CM and surfactant-enhanced NAA penetration will be discussed, in relation to diffusion in the non-living, non-metabolic plant cuticle.
Foliar application of plant growth regulators (PGR) is an established horticultural practice. We are using a finite dose system to examine diffusion of 14C-labeled PGRs, primarily napththaleneacetic acid (NAA), from aqueous droplets and deposits through enzymatically isolated plant cuticles (CM) as affected by spray adjuvant chemistry, solution pH, and epicuticular wax. Recent studies have focused on a nonbuffered aqueous medium, which approximates field application conditions. Despite the negligible buffering capacity of the spray solution, there were significant differences in NAA diffusion with solution pH. At pH 3.2, NAA (pKa = 4.2) diffusion was two-fold greater than at pH 5.2. Additives (surfactants, urea, and urea:NH4NO3, 1:1 mixture) in the spray solutions increased the initial rate and absolute amount of NAA diffused. The polyethoxalated octylphenol surfactant (Triton-X) TX-45 (EO 5.5) enhanced rate and quantity of NAA diffusion. This enhancement was observed with CM, but not after removal of the epicuticular waxes, implicating an interaction between surfactant and waxes. Urea, over a four-fold concentration range, increased NAA diffusion 5% to 31% after 144 h. The urea:NH4NO3 mixture increased NAA diffusion to a greater extent at pH 5.2 (+136%) than at pH 3.2 (+8.4%) after 144 h.
Morphological and physical characteristics of the cuticular membrane (CM) of selected cultivars of sweet cherry (Prunus avium L.) fruit were studied relative to rain-induced cracking. Two characteristics of the CM may be determinants in rain-induced fruit cracking. The surface morphology and chemistry determine surface wettability and water retention, and the morphology and physicochemical characteristics its water permeability. The fruit epidermis as well as the guard cell walls adjacent to the outer vestibule and stomatal pore are covered by a thin lipoidal CM. Stomata were present at a frequency of 0.1 to 2 per mm2 depending on cultivar and fruit surface position. However, most appeared nonfunctional with many pores partially or completely occluded with wax-like material. There was no evidence of water (containing fluorescein or AgNO3) penetration into stomatal pores following surface application or submerging fruit for short periods. There was stomatal pore penetration when submerged fruit were infiltrated by reduced pressure in the presence of 0.1% L-77. Preferential sorption of AgNO3 and fluorescein by cuticular ledges and guard cells was noted. The epicuticular wax (ECW) had no significant fine-structure. The CM was isolated enzymatically (cellulase/pectinase) and found to be 1 to 2 μm thick with an area weight of 1.2 to 2.3 g·m–2, of which 25% to 40% was chloroform/methanol (1: 1by vol.) soluble. Fractionation of the chloroform/methanol fraction indicated the presence of four groups of nonpolar constituents. The fruit surface was moderately difficult to wet, forming contact angles of 85% to 105%, and with an estimated critical surface tension in the range of 16-24 mN·m–1. Fruit water loss (transpiration) and uptake on submersion was followed and found to be complex. Transpiration increased with an increase in temperature, and both rate of transpiration and water uptake increased after removal of the epicuticular and cuticular waxes. Pathways of water uptake and the significance of our findings to rain-induced fruit cracking will be discussed.
NAA, a weak organic acid plant growth regulator (pKa 4.2), penetrates the plant cuticle preferentially as an undissociated molecule (<10% dissociated at pH 3.2). We have reported, using a finite dose diffusion system, that NH4NO3 (AN, 8 mM) at pH 5.2 (>90% dissociated) enhanced the penetration of 14C-NAA through isolated tomato fruit cuticular membranes (CM). AN appears to preferentially enhance penetration of the dissociated NAA molecule over the nondissociated form. A possible mode of action is that AN affects the cuticle matrix, allowing for greater NAA penetration. Acid treatment (4 N HCl) of the cuticle, which alters the cuticle's ionic characteristics, resulted in a 10% reduction in NAA penetration from droplets in the presence of AN. When AN (80 mM) was included in the receiver solution of the diffusion cell in an effort to infuse the cuticle matrix, NAA penetration was not increased compared to when AN was present in the applied droplet. AN (8 mM) also increased NAA penetration through dewaxed tomato cuticular membranes (DCM; +252% for DCM vs. +190% for CM in 120 h). Since AN only enhances NAA penetration when included with the NAA in the treatment droplet, the AN effect may be related to the role of the droplet/deposit (droplet residue) as a donor. This conclusion is further supported with sorption studies, where AN over a 100-fold concentration range (0.8–80 mM) did not increase NAA sorption by tomato fruit CM and where no deposit is present. The role of the physicochemical nature of the deposit, including the chemical/ionic characteristics of any additive (i.e., AN) and active ingredient will be discussed.
The cuticle is the prime barrier to penetration of foliar applied plant growth regulators, which must penetrate and be transported to a reaction site before a response can be induced. Urea has enhanced performance of Fe and Zn foliar sprays and a mixture of urea and ammonium nitrate (WAN) the performance of some herbicides. The mechanism of this enhancement is not clear. We find that urea and UAN increased 14C-NAA transport across enzymatically isolated tomato fruit cuticular membranes (CM) from simulated spray droplets using a finitedose diffusion system. The initial rate and total amount of NAA penetrated was significantly increased relative to NAA alone, the enhancement being greater for UAN than urea (total amount 101% vs. 78% at 120 hours) and for the NAA anion (pH 5.2, pKa 4.2) than for the nondissociated (pH 3.2) moiety. When evaluating the concentration effect of urea and NH4NO3 individually, the greatest enhancement with urea was at 62 mm and with NH4NO3 at 8 mm. Generally the effect of urea was significantly less than NH4NO3 (+24% vs. 296%). NAA penetration was greater with NH4NO3 than with KNO3 or Ca(NO3)2 or when the nitrate anion was replaced with sulfate or phosphate. Transcuticular penetration of NAA was enhanced greatly (190% in 120 hours) on removal of cuticular waxes; however, penetration was further increased (252% in 120 hours) by adding 8 mm NH4NO3. Methylamine hydrochloride (CH3NH2.HC1, 8 mm) also increased NAA diffusion, the initial slopes (>8 hours) were 23, 14, and 2 pmols·h–1 for methylamine, ammonium nitrate, and NAA alone, respectively, while the percent of applied that penetrated after 120 hours was 68.5, 67.6, and 21.4 for methylamine, ammonium nitrate, and NAA alone, respectively. The enhancement of NAA penetration by NH4NO3 equaled or exceeded that obtained with a group of surfactants of diverse chemistries. When the surfactant Triton X-100 was compared with NH4NO3, initial penetration was more rapid with ammonium nitrate (11.7 vs. 7.3 pmols·h–1) but percent penetrating after 120 hours was greater for Triton X-100 (80.5 vs. 66.8). The possible action of NH4NO3 on NAA uptake will be discussed.