Plant Growth Regulators Gibberellic Acid Research Articles

Interaction of Gibberellic Acid and Glyphosate on Growth and Phenolic Metabolism in Soybean Seedlings

The plant growth regulator gibberellic acid (GA) and the herbicide glyphosate were examined for their possible interactions with growth and phenolic metabolism in soybean [Glycine max (L.) Merr. Cv. Hill] seedlings. GA caused increases in phenylalanine ammonia-lyase activity (PAL) (per axis basis) above those of the control seedling levels 48 h after treatment in light-grown seedlings. This effect increased to two-fold greater than control levels by 72 and 96 h after treatment. In dark-grown plants, GA had no effect on PAL levels at 24 h, reduced levels at 48 and 72 h, and increased PAL at 96 h. Early studies in our lab reported that glyphosate increased PAL levels, and also reduced hydroxyphenolic compound accumulation in both light- and dark-grown soybean seedlings. Treatments of GA plus glyphosate caused additive increases in PAL activity in light-grown seedlings, but GA lowered glyphosate’s increase in PAL levels at 48–96 h after treatment in dark-grown seedlings. GA had little effect on hydroxyphenolic compound levels in either light- or dark-grown seedlings. GA treatment alone did not significantly affect root elongation, but stimulated hypocotyl and epicotyl elongation and caused marginal reversal of glyphosate inhibition of elongation in roots, hypocotyls, and epicotyls in light-grown plants. These results show some differential effects of GA and glyphosate on growth and phenolic metabolism, and their interactions that are dependent on plants grown in light or darkness.

Effect of Plant Growth Regulators Giberellic acid (GA3) And Salicylic Acid (SA) on Growth and Yield of Carnation (Dianthus caryophyllus) under Naturally Ventilated Polyhouse

The present investigation entitled “Effect of plant growth regulators Gibberellic Acid (GA3) and Salicylic Acid (SA) on the growth and yield of carnation under naturally ventilated polyhouse”, was carried out during November, 2021 to April 2022 in, Naturally Ventilated Polyhouse, Department of Horticulture, Naini Agricultural Institute, SHUATS, it was concluded that the plant growth regulators treatments rendered their significant effect on almost all the growth, flowering and yield characters as well as quality of carnation. The treatment T3 i.e. application of GA3 @ 200 ppm was found superior in terms of Plant height (102.27cm), number of leaves (203.67), number of shoots (7.67), internodal length (7.73cm), number of internodes per shoot (17), bud length (10cm), bud diameter (2.87cm), flower diameter (6.80cm), flower length (5.67cm), flower stalk length (96.93cm), vase life (10.33), number of cut flower stalks per plant (7.00) and number of cut flower stalks per square meter (175). However, plants treated with SA @ 120ppm took minimum days for bud initiation (112.67 days) and the days taken to bud opening (24.67). Among the different treatments the highest Gross return (Rs/200m2) (7,00,000), Net return (441565), benefit cost ratio (2.70) was obtainedunder the use of GA3 @ 200 ppm (T3).

Effects of Exogenous Gibberellic Acid in Huanglongbing-affected Sweet Orange Trees under Florida Conditions—II. Fruit Production and Tree Health

Fruit production of sweet orange (Citrus sinensis) in Florida has been declining with the presence of Huanglongbing [HLB; Candidatus Liberibacter asiaticus (CLas)] disease. Through disruption of the balance of endogenous hormone levels, the disease has negative impacts on fruit development, mature fruit retention, and overall tree health. Thus, the goal of this research was to determine whether plant growth regulator gibberellic acid (GA3) can be used to improve the production issues caused by HLB. For ‘Valencia’ sweet orange, although foliar applied GA3 from September to January (33 mg·L−1 for five applications) resulted in 50% decrease in bloom the following spring (results presented in Tang et al., 2021), this treatment did not cause reduction in yield of current and subsequent crops. Moreover, a 30% average increase in yield in GA3-treated trees was observed over a period of 4 years. The size of mature fruit was also increased (by 4% to 5%) with reduced fruit drop rate near harvest in GA3-treated trees compared with nontreated control trees. Furthermore, the canopy density, an indicator of HLB severity, was maintained in trees applied with GA3 (from 90.8% light interception to 90.4%). In contrast, there was a substantial decrease in canopy density for control trees (from 91.6% to 84.0%). Gene expression analysis of abscission zone and leaves indicated that GA3-treated trees had enhanced oxidative stress mitigation mechanism and plant defense response. Given that there is no cure for HLB, these results presented a possible remedy of using GA3 in sustaining tree health for field-grown sweet orange affected by HLB.

Gibberellic acid-induced hepatorenal dysfunction and oxidative stress: Mitigation by quercetin through modulation of antioxidant, anti-inflammatory, and antiapoptotic activities.

The plant growth regulator gibberellic acid (GA3) is widely used in agriculture in many countries. However, little is known about its danger to human health or its physiologic and biochemical pathways. Our study examined the effect of GA3 on liver and kidney function and the effect of quercetin on the hepatorenal toxicity induced by GA3 in four groups of male albino rats. For 4weeks, the control group (CNT) received saline, the quercetin group (QR) received daily intraperitoneal injections of quercetin (50mg/kg/BW) dissolved in saline, the gibberellic acid group (GA3) received GA3 (55mg/kg/BW) via oral gavage, and the protective group (QR) was injected with quercetin and gavaged with GA3 in the same doses used in the QR and GA3 groups (50mg/kg/BW +GA3 and 55mg/kg/BW). GA3 induced liver and kidney injury, as shown by elevated serum glutamic pyruvic transaminase, glutamic oxaloacetic transaminase,and gamma-glutamyl transferase (GPT, GOT, and GGT) as well as increased levels of creatinine, urea, and uric acid. Hepatorenal toxicity was demonstrated by a significant increase in levels of serum and tissue malondialdehyde (MDA) and decreased antioxidant enzyme activity, such as catalase (CAT) and superoxide dismutase (SOD), accompanied by a subsequent decrease in glutathione peroxidase (GPx) levels in liver and kidney tissue of GA3-treated rats. Administration of quercetin (QR) significantly protected hepatorenal tissue against the toxic effect of GA3 through normalization of the hepatic and renal function markers. It also retrieved the antioxidant ability by modulating the hepatorenal toxic effect at the molecular level through upregulation of antiapoptotic genes and downregulation of transforming growth factor-β1 (TFG-β1), cyclooxygenase-2 (COX-2), and nuclear factor-kappa B (NF-κB). Impairment of liver and kidney function was confirmed by histologic and immunohistochemical analyses. Pretreatment with quercetin was effective at attenuating histopathologic changes in hepatic and renal tissues by regulating the immunoexpression of caspase-3 and Bcl-2 to return them to more normal values. PRACTICAL APPLICATIONS: The confirmed hepatorenal dysfunction caused by GA3 was ameliorated by quercetin administration. Moreover, quercetin demonstrated the potential to reverse hepatorenal dysfunction by regulating inflammatory and antioxidant properties, inhibiting the production of free radicals and inflammation-associated cytokines, and modulating antioxidants and antiapoptotic activity.

Effects of the nitrification inhibitor nitrapyrin and the plant growth regulator gibberellic acid on yield-scale nitrous oxide emission in maize fields under hot climatic conditions

Nitrification inhibitors are widely used in agriculture to mitigate nitrous oxide (N2O) emission and increase crop yield. However, no concrete information on their mitigation of N2O emission is available under soil and environmental conditions as in Pakistan. A field experiment was established using a silt clay loam soil from Peshawar, Pakistan, to study the effect of urea applied in combination with a nitrification inhibitor, nitrapyrin (2-chloro-6-tri-chloromethyl pyridine), and/or a plant growth regulator, gibberellic acid (GA3), on N2O emission and the nitrogen (N) uptake efficiency of maize. The experimental design was a randomized complete block with five treatments in four replicates: control with no N (CK), urea (200 kg N ha–1) alone, urea in combination with nitrapyrin (700 g ha–1), urea in combination with GA3 (60 g ha–1), and urea in combination with nitrapyrin and GA3. The N2O emission, yield, N response efficiency, and total N uptake were measured during the experimental period. The treatment with urea and nitrapyrin reduced total N2O emission by 39%–43% and decreased yield-scaled N2O emission by 47%–52%, relative to the treatment with urea alone. The maize plant biomass, grain yield, and total N uptake increased significantly by 23%, 17%, and 15%, respectively, in the treatment with urea and nitrapyrin, relative to the treatment with urea alone, which was possibly due to N saving, lower N loss, and increased N uptake in the form of ammonium; they were further enhanced in the treatment with urea, nitrapyrin, and GA3 by 27%, 36%, and 25%, respectively, probably because of the stimulating effect of GA3 on plant growth and development and the reduction in biotic and abiotic stresses. These results suggest that applying urea in combination with nitrapyrin and GA3 has the potential to mitigate N2O emission, improve N response efficiency, and increase maize yield.

Effect of gibberellin, methyl jasmonate and myoinositol on biomass and eicosapentaenoic acid productivities in the eustigmatophyte Monodopsis subterranea CCALA 830

Omega-3 fatty acids sourced from microalgae are in demand due to their therapeutic potential and sustainability of the source. The present study was focused towards enhancing eicosapentaenoic acid (EPA) productivity in the marine Chrysophyte Monodopsis subterranea through media supplementation using the plant growth regulators gibberellic acid, methyl jasmonate and myoinositol. The effect of three concentrations (1 μM, 10 μM and 100 μM) of supplements on biomass yield, fatty acid composition, total fatty acid (TFA) and EPA productivity were analyzed at three phases of microalgal growth. Supplementation with 100 μM GA increased biomass yield and TFA productivity by 3.3-fold and 3.9-fold respectively at the mid-exponential phase and EPA productivity increased up to 3.2-fold at the late-exponential phase. The addition of 1 μM MeJA stimulated biomass accumulation to 2.7-fold at the late-stationary phase, but this effect was reduced to 1.1–1.4-fold at higher concentrations. The addition of 10 μM MI also showed a positive effect on TFA productivity and thus increased EPA productivity by 1.8-fold at the late-stationary phase. Supplementation with 100 μM GA, 100 μM MeJA and 10 μM MI at the late-stationary phase increased all fatty acids except 18:0 and 18:2. EPA was negatively correlated with 18:2 at all three growth phases. Principal components analysis indicated a strong relationship between key fatty acids and EPA at all three growth stages and associated with 100 μM GA.

Influence of plant growth regulators on growth, yield, and quality of tomato grown under high temperature in the tropics in the summer

ABSTRACT Yield of tomato (Solanum lycopersicum L.) is affected by high day and night temperature and high humidity which can occur together. Plant growth regulators may minimize adverse environmental effects to increase tomato yield. A field experiment was conducted in 2015 and 2016 to evaluate effects of foliar-applied plant growth regulators gibberellic acid (GA3) and 4-chloro phenoxy acetic acid (4-CPA) on growth, yield, and quality of tomato subjected to high day and night temperature (around 30°C) and high humidity, avg. 83%. Concentrations of GA3 were 0, 25, 50, 100 or 125 mg GA3∙L−1, and concentrations of 4-CPA were 0 or 20 mg 4-CPA∙L−1. Gibberellic acid was applied alone during pre-anthesis at 30 days after transplanting; 4-CPA was applied at anthesis and a second dose 7 days later. Application of 50 mg GA3∙L−1and 20 mg 4-CPA∙L−1 produced the heaviest individual fruit (76.2 g) and greatest yield (46.1 t∙ha−1) with higher total soluble solids (4.48 °Brix) and vitamin C (31.6 mg/100 g). Combined application of 50 mg GA3∙L−1 and 20 mg 4-CPA.L−1 was economical and could be applied under high temperature and humidity to increase quantity and quality of tomato fruit.

Potential Toxicity of Plant Growth Regulator Gibberellic acid (GA3) on the Pancreatic Structures and Functions in the albino rat

Potential Toxicity of Plant Growth Regulator Gibberellic acid (GA3) on the Pancreatic Structures and Functions in the albino rat

γ-Polyglutamic acid/chitosan nanoparticles for the plant growth regulator gibberellic acid: Characterization and evaluation of biological activity

The growth regulator gibberellic acid (GA3) has several uses in the field, improving germination, plant development, productivity, and the quality of food. This work describes the development of a nanocarrier system for GA3, based on the poly(γ-glutamic acid) (γ-PGA) and chitosan (CS) polymers, Nanoparticles without GA3 (nano-γPGA/CS-GA3) showed colloidal characteristics, with an average size of 117±9nm, PDI of 0.43±0.07, and zeta potential of −29±0.5mV. The encapsulated nanoparticles (nano-γPGA/CS-GA3) presented an average size of 134±9nm, PDI of 0.35±0.05, zeta potential of 27.9±0.5mV, and 61% encapsulation. The images of nanoparticles observed by Transmission and scanning electron microscopy (TEM and SEM) showed a spherical shape of the nanoparticles. The system showed sustained release, with 58% release after 48h. Evaluation of thermal properties using DSC and TGA analyses indicated that there was an interaction between the CS and γ-PGA polymers. In tests using Phaseolus vulgaris seeds, nano-γPGA/CS-GA3 showed high biological activity, enhancing the rate of germination in the first day (50–70%) when compared with free GA3 (10–16%). Encapsulated GA3 was also more efficient than the free hormone in the increase of leaf area and the induction of root development (including the formation of lateral roots). These effects were not observed when seeds were treated with nano-γPGA/CS without GA3. The results demonstrated the considerable potential of nano-γPGA/CS-GA3 for use in agriculture.

Chitosan nanoparticles as carrier systems for the plant growth hormone gibberellic acid

This work concerns the development of nanocarriers composed of alginate/chitosan (ALG/CS) and chitosan/tripolyphosphate (CS/TPP) for the plant growth regulator gibberellic acid (GA3). ALG/CS nanoparticles with and without GA3 presented mean size of 450±10nm, polydispersity index (PDI) of 0.3, zeta potential of −29±0.5mV, concentrations of 1.52×1011 and 1.92×1011 nanoparticles mL−1, respectively, and 100% encapsulation efficiency. CS/TPP nanoparticles with and without GA3 presented mean size of 195±1nm, PDI of 0.3, zeta potential of +27±3mV, concentrations of 1.92×1012 and 3.54×1012 nanoparticles mL−1, respectively, and 90% encapsulation efficiency. The nanoparticles were stable during 60days and the two systems differed in terms of the release mechanism, with the release depending on factors such as pH and temperature. Bioactivity assays using Phaseolus vulgaris showed that the ALG/CS-GA3 nanoparticles were most effective in increasing leaf area and the levels of chlorophylls and carotenoids. The systems developed showed good potential, providing greater stability and efficiency of this plant hormone in agricultural applications.

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Fatty acid esters produced by Lasiodiplodia theobromae function as growth regulators in tobacco seedlings

The Botryosphaeriaceae are a family of trunk disease fungi that cause dieback and death of various plant hosts. This work sought to characterize fatty acid derivatives in a highly virulent member of this family, Lasiodiplodia theobromae. Nuclear magnetic resonance and gas chromatography-mass spectrometry of an isolated compound revealed (Z, Z)-9,12-ethyl octadecadienoate, (trivial name ethyl linoleate), as one of the most abundant fatty acid esters produced by L. theobromae. A variety of naturally produced esters of fatty acids were identified in Botryosphaeriaceae. In comparison, the production of fatty acid esters in the soil-borne tomato pathogen Fusarium oxysporum, and the non-phytopathogenic fungus Trichoderma asperellum was found to be limited. Ethyl linoleate, ethyl hexadecanoate (trivial name ethyl palmitate), and ethyl octadecanoate, (trivial name ethyl stearate), significantly inhibited tobacco seed germination and altered seedling leaf growth patterns and morphology at the highest concentration (0.2 mg/mL) tested, while ethyl linoleate and ethyl stearate significantly enhanced growth at low concentrations, with both still inducing growth at 98 ng/mL. This work provides new insights into the role of naturally esterified fatty acids from L. theobromae as plant growth regulators with similar activity to the well-known plant growth regulator gibberellic acid.

Determination of Gibberellic Acid (GA3)-Induced Oxidative Stress in a Model Organism Galleria mellonella L. (Lepidoptera: Pyralidae).

The plant growth regulator gibberellic acid (GA3) is known to negatively impact growth and development of insects. In this study, larvae of Galleria mellonella L. (Lepidoptera: Pyralidae) were fed a diet with varying dosages of GA3 to investigate how antioxidant enzymes are influenced. Activity levels in last instars reared in laboratory at 25 ± 2°C, 60 ± 5% relative humidity, and a photoperiod of 12:12 (L:D) h were measured for superoxide dismutase (SOD), glutathione S-transferase (GST), and catalase (CAT). Treatment with GA3 in diet resulted in a remarkable increase in the activities of both SOD and GST at lower GA3 doses (50-1,000 ppm) with respect to control and higher doses. The activity of CAT in the hemolymph of last instars significantly increased at all doses when compared with that in the hemolymph of untreated larvae. This trend in the increase of CAT was not dose-wise, except for the significant increases at 2,000 and 5,000 ppm when compared with that of untreated and all treated groups. Consequently, our results showed that GA3 is effective at activating the antioxidant defense system of insects as a source of free radical and can be toxic for larvae in a dose-dependent manner. Therefore, we suggest that the increase in the activity of GST, SOD, and CAT in larvae may indicate a physiological adaptability to compensate for GA3-induced stress.

Effect of gibberellic acid (GA), indole acetic acid (IAA) and benzylaminopurine (BAP) on the synthesis of essential oils and the isomerization of methyl chavicol and trans-anethole in Ocimum gratissimum L.

Basil (O. gratissimum L) is a aromatic and medicinal plant widely used in traditional medicine in Morocco. The aim of this work was to study the effect of three plant growth regulators gibberellic acid (GA), indole 3-acetic acid (IAA) and benzylaminopurine (BAP) on the content and composition of essential oils of this plant, especially on the main compound (methyl chavicol) and its isomer (the trans-anethole).The results showed a wide variation on yield, content and range of the molecule constituent of oil, with a balance of appearances and/or disappearances of a few molecules.GA caused a slight decrease in the oil yield (0.2%), but it increased the diversity of compounds (17 molecules) with the appearance of four new compounds (naphthalene, camphor, germacrene-D, and ledene) and disappearance of (β cedrene, azulene). This variation also caused a very important decrease in the main compound (methyl chavicol) and increases its isomer (trans-anethole).IAA and BAP caused an increase in the yield of essential oil (0.30% and 0.32% respectively) without much influence on the main compounds, but with some change in the composition such as the appearance of (germacrene-D) and the disappearance of (aristolene).

Effects of Gibberellic Acid on Hemocytes ofGalleria mellonellaL. (Lepidoptera: Pyralidae)

The impacts of different doses of the plant growth regulator gibberellic acid (GA(3)) in diet on the number of total and differential hemocytes, frequency of apoptotic, and necrotic hemocytes, mitotic indices, encapsulation, and melanization responses were investigated using the greater wax moth Galleria mellonella L. (Lepidoptera: Pyralidae) larvae. Total hemocyte counts increased in G. mellonella larvae at all treatment doses whereas GA(3) application had no effect on the number of different hemocyte types. The occurrence of apoptosis, necrosis and mitotic indices in GA(3) treated and untreated last instars were detected by acridine orange or ethidium bromide double staining by fluorescence microscopy. While the ratio of necrotic hemocytes increased at all GA(3) treatments, that of late apoptotic cells was only higher at doses >200 ppm when compared with untreated larvae. The percentage of mitotic index also increased at 5,000 ppm. Positively charged DEAE Sephadex A-25 beads were used for analysis of the levels of encapsulation and melanization in GA(3) treated G. mellonella larvae. At four and 24 h posttreatments with Sephadex A-25 bead injection, insects were dissected under a stereomicroscope. Encapsulation rates of larval hemocytes were dependent on the extent of encapsulation and time but not treatment groups. While the extent of melanization of hemocytes showed differences related to time, in general, a decrease was observed at all doses of GA(3) treated larvae at 24 h. We suggest that GA(3) treatment negatively affects hemocyte physiology and cell immune responses inducing cells to die by necrosis and apoptosis in G. mellonella larvae.

EFFECT OF GIBBERELLIC ACID ON THE PERSISTENCE OF CERTAIN PESTICIDES ON/IN GRAPE FRUITS

Recommended application was done by spray regime for two pesticides, namely diniconazole, fenitrothion and a plant growth regulator (gibberellic acid) and their mixtures. Their residues were determined after different successive treatments in and on grape fruits in two consecutive years. The initial deposits of the fungicide diniconazole when used alone were 0.49 and 0.50 ppm for 2006 and 2007, respectively. While the initial deposits of fenitrothion reached to 11.35 and 11.19 ppm for the same interval, respectively. Initial deposits of the plant growth regulator gibberellic acid were 30.52 and 30.42 ppm for the same period, respectively. After mixing the tested pesticides and the plant growth regulator, the initial deposits loss reached 38.77 and 42.0 % for diniconazole, 43.08 and 48.16 % for fenitrothion and 6.88 and 7.00 % for gibberellic acid for the same seasons, respectively. A significant degradation was recorded with the mixture of the two tested pesticides and plant growth regulator compared with that occurred when pesticide was used alone at the two studied seasons.

Efeito da aplicação de ácido giberélico no crescimento da palmeira-ráfia

A palmeira-ráfia possui grande importância econômica devido ao seu interesse comercial. É uma planta ornamental muito apreciada por sua beleza e utilização em ambiente interno, crescendo bem em locais com sombra e apresentando resistência à exposição solar. O valor econômico dessa planta está relacionado à altura e número de hastes. O objetivo do trabalho foi promover o crescimento da planta, através da aplicação do regulador de crescimento ácido giberélico (GA3). Mudas de palmeira-ráfia com aproximadamente um ano de idade foram aspergidas a cada 21 dias (quatro aplicações) com solução de ácido giberélico nas concentrações de 0; 75; 150; 225; e 300 mg.L-1. Os resultados indicaram que a aplicação do ácido giberélico foi eficiente na promoção do crescimento da espécie, sendo esse incremento significativo nas variáveis: comprimento dos pecíolos, lâminas foliares e altura da planta. Não se observou diferença no acúmulo de matéria seca ou fresca do pecíolo entre os tratamentos, e plantas do tratamento-controle foram superiores às dos tratamentos 225 e 300 mg L-1, nas variáveis diâmetro do colo e massa fresca e seca das raízes. Devido ao incremento na altura das mudas e à alteração da disposição das folhas da palmeira, a utilização do ácido giberélico pode ser recomendada para a produção de mudas de alta qualidade.

Apoptosis in barley aleurone during germination and its inhibition by abscisic acid.

During germination of barley grains, DNA fragmentation was observed in the aleurone. The appearance of DNA fragmentation in the aleurone layer, observed by TUNEL staining in aleurone sections, started near the embryo and extended to the aleurone cells far from the embryo in a time dependent manner. The same spatial temporal activities of hydrolytic enzymes such as alpha-amylase were observed in aleurone. DNA fragmentation could also be seen in vitro under osmotic stress, in isolated aleurone. During aleurone protoplast isolation, a very enhanced and strong DNA fragmentation occurred which was not seen in protoplast preparations of tobacco leaves. ABA was found to inhibit DNA fragmentation occurring in barley aleurone under osmotic stress condition and during protoplast isolation, while the plant growth regulator gibberellic acid counteracted the effect of ABA. Addition of auxin or cytokinin had no significant effect on DNA fragmentation in these cells. To study the role of phosphorylation in ABA signal transduction leading to control of DNA fragmentation (apoptosis), the effects of the phosphatase inhibitor okadaic acid and of phenylarisine oxide on apoptosis were studied. We hypothesize that the regulation of DNA fragmentation in aleurone plays a very important role in spatial and temporal control of aleurone activities during germination. The possible signal transduction pathway of ABA leading to the regulation of DNA fragmentation is discussed.

Investigation of Effects of Plant Growth Regulators on Liposome Fluidity and Permeability

AbstractNaturally occurring plant growth regulators gibberellic acid (GA), indole‐3‐acetic acid (IAA), abscisic acid (ABA), ethylene and other growth regulating compounds such as 5‐methyl‐7‐chloro‐4‐ethoxycarbonylmethoxy‐2,1,3‐benzothiadiazole (TH) and 2,4 dichlorophenoxyacetic acid (2,4‐D), had no effect on the partition behavior of a piperidine based spin label in liposomes composed of pure or mixed phospholipids or a phospholipid‐sterol mixture. Although no effect on fluidity was observed, TH significantly increased the initial rate of swelling of soybean lecithin‐sitosterol liposomes in isotonic glycerol. IAA and ethylene did not influence this rate but ABA, GA and 2,4‐D inhibited the initial rate of swelling. Lipid composition of liposomes determined the extent and direction of the effects on swelling rates. The observed swelling behavior was, therefore, not related to fluidity of the bulk membrane lipids but was due, instead, to modification of the access of glycerol to the phospholipid bilayer surface or, alternatively, to the creation of polar channels into the liposomes.