Leaf Extension in Zea mays: II. LEAF EXTENSION IN RESPONSE TO INDEPENDENT VARIATION OF THE TEMPERATURE OF THE APICAL MERISTEM, OF THE AIR AROUND THE LEAVES, AND OF THE ROOT-ZONE

The temperatures of the roots, the apical meristem, and theshoots of Zea mays plants were varied independently of eachother and the rates of leaf extension were measured. When thetemperature of the apical meristem and region of cell expansionat the base of the leaf was kept at 25 °C, changes of leafextension in response to changes of root and shoot temperatureswere less pronounced. When the temperature of the meristematicregion was changed by increments of 5 or 10 °C from 0 to40 °C, and the root and shoot temperatures were kept at25 °C, rapid changes in leaf extension occurred. It was concluded that the rates of leaf extension were controlledat root-zone temperatures of 5 to 35 °C by heating or coolingof the meristematic region. Changes in rates of leaf extensionin response to changes in air temperature were attributed todirect effects on the temperature of the meristematic regionand on the physiology of the leaf.     

Low Temperature Effects on Early Vegetative Growth, Leaf Gas Exchange and Water Potential of Chilling-sensitive and Chilling-tolerant Crop Species

Two chilling-sensitive (Phaseolus vulgaris L., Zea mays L.)and two chilling-tolerant (Pisum sativum L., Spinacia oleraceaL.) species were raised in growth chambers under warm (28/18°Cday/night cycle) and cool (18/12°C) temperature regimes.Growth analysis techniques were used to evaluate leaf area andbiomass partitioning during early autotrophic growth. Plantsacclimated to both temperatures were measured for leaf gas exchangeand water potential (     

Interrelationships between the Nature of the Light Source, Ambient Air Temperature, and the Vegetative Growth of Different Species within Growth Cabinets

The interacting effects within growth cabinets of the natureof the light source and the temperature of the ambient air on14 parameters of vegetative growth have been examined for Gotsypiumhirsutum, Helianthus annuus, Phaseolus vulgaris, and Zea mays.The comparisons were between fluorescent lighting and mixtureswith tungsten lamps (17 and 26 per cent of the total wattage),adjusted to give 32 400 lx and covering six equally spaced andconstant air temperatures from 10 to 35 °C. The relationships are highly complex. Species differ in theirresponses to the three light sources. Moreover, for some parameters,but only for some species, there are highly significant interactionsbetween the nature of the light source and the temperature level.Part of this complexity is due to the varying growth potentialof species at low and high temperatures but important contributingfactors are the differences between the temperatures of theambient air and the plant parts. Such disparities may be positiveor negative and are greater for leaf as against root temperature.The divergencies are larger during the day and largest for leaftemperature. With the higher proportion of tungsten lamps theleaf day temperature ranges from 7.1 °C above an ambienttemperature of 10 °C to 2.5 °C above 35 °C ambient.For fluorescent lighting the leaves are 2.8 °C above ambientat 10 °C and 2.5 °C below at 35 °C. When variations in the response are examined against the meaninternal day temperature of the plant, for some parameters,but not others, differences between light sources are no longerevident. Here again the reactions of species differ. It is concluded firstly that if only ambient air temperaturesare recorded errors will arise in the interpretation of theinduced biological responses, and secondly that departures fromthe ambient air temperatures will be dependent on the designof the cabinet and the lighting system.     

Effect of Root and Shoot Meristem Temperature on Shoot to Root Dry Matter Partitioning and the Internal Concentrations of Nitrogen and Carbohydrates in Maize and Wheat

Maize (Zea mays L.) and spring wheat (Triticum aestivum L.)were grown in nutrient solution at uniformly high air temperature(20 °C), but different root zone temperatures (RZT 20, 16,12 °C). To manipulate the ratio of shoot activity to rootactivity, the plants were grown with their shoot base includingthe apical meristem either above (i.e. at 20 °C) or withinthe nutrient solution (i.e. at 20, 16 or 12 °C). In wheat, the ratio of shoot:root dry matter partitioning decreasedat low RZT, whereas the opposite was true for maize. In bothspecies, dry matter partitioning to the shoot was one-sidedlyincreased when the shoot base temperature, and thus shoot activity,were increased at low RZT. The concentrations of non-structuralcarbohydrates (NSC) in the shoots and roots were higher at lowin comparison to high RZT in both species, irrespective of theshoot base temperature. The concentrations of nitrogen (N) inthe shoot and root fresh matter also increased at low RZT withthe exception of maize grown at 12 °C RZT and 20 °Cshoot base temperature. The ratio of NSC:N was increased inboth species at low RZT. However this ratio was negatively correlatedwith the ratio of shoot:root dry matter partitioning in wheat,but positively correlated in maize. It is suggested that dry matter partitioning between shoot androots at low RZT is not causally related to the internal nitrogenor carbohydrate status of the plants. Furthermore, balancedactivity between shoot and roots is maintained by adaptationsin specific shoot and root activity, rather than by an alteredratio of biomass allocation between shoot and roots.Copyright1994, 1999 Academic Press Wheat, Triticum aestivum, maize, Zea mays, root temperature, shoot meristem temperature, biomass allocation, shoot:root ratio, carbohydrate status, nitrogen status, functional equilibrium     

The Growth and Development of Maize (Zea mays L.) at Five Temperatures

The objectives of this work were to measure growth and developmentrates over a range of temperatures and to identify processeswhich may limit vegetative yield of maize (Zea mays L.). Twosingle cross Corn Belt Dent maize hybrids were grown from sowingin a diurnal temperature regime of 16/6 °C day/night andin constant temperature environments of 16, 20, 24 and 28 °C.The 16/6 °C environment was close to the minimum for sustainedgrowth and 28 °C was near the optimum. Entire plants wereharvested at stages with 4, 6, 7 and 8 mature leaves in alltemperature treatments except 20 °C in which the final twoharvests were carried out at 9 and 10 mature leaves. Mean totalleaf number varied between 19.5 and 16.0 with the maximum occurringat 16/6 °C. Although harvests were carried out at comparableleaf numbers, and hence at similar developmental stages, thetime interval between sowing and harvest decreased considerablyas temperatures increased. The relative rates of dry weight and leaf area accumulationwith time increased with a Q₁₀ of 2.4 between 16 and 28 °C,while leaf appearance rate increased with a Q₁₀ of 2.9 overthe same range; both rates were highest at 28 °C. Althoughdry matter partitioning to the shoots increased with temperature,the area of individual leaves varied in a systematic patternwhich resulted in maximum leaf area, leaf area duration andconsequently dry weight being realized at 20 °C for anygiven stage of development. Zea mays, corn, low temperature stress, temperature response, growth, development     

Internal Root Anatomy of Maize Seedlings (Zea mays L.) as Influenced by Temperature and Genotype

The objectives of this investigation were to determine: (a)the general effect of temperature on internal root anatomy;(b) whether genotypic differences in such root traits exist;and (c) the association between internal root traits and shootgrowth, lateral root branching and cold tolerance of maize (Zeamays L.). Seedlings of 20 central European hybrids were grownunder high or low temperature (25/22·5 °C or 15/12·5°C day/night temperatures) until the third leaf was fullyexpanded. Light microscopy of cross sections revealed a largerdiameter of primary roots at low temperature which was due toa larger stele diameter and a thickening of the cortex. Concurrently,an increase in total cross sectional area of metaxylem elementswas obtained. It is assumed that the modification of the internalroot structure by temperature has an effect on both axial andradial water flow capacity. For all anatomical traits studied,variability between genotypes was apparent under both growingconditions. Furthermore, different genotypic responses to temperaturewere observed. However, basic differences between cold-tolerantand cold-sensitive genotypes did not exist. While at high temperatureroot traits and shoot growth were significantly and positivelycorrelated, at low temperature the correlation coefficient wasinsignificant. Consequently, it was not possible to characterizethe performance of the shoot at low temperature based on anatomicaltraits of the root. Moderate, positive correlation coefficientswere obtained between internal root traits and lateral rootbranching. The potential use of root anatomical traits as indirectselection criteria is discussed. Chilling tolerance, genotypes, root anatomy, Zea mays L     

Influences of Water Status, Temperature, and Root Age on Daily Patterns of Root Respiration for Two Cactus Species

Daily patterns of root respiration measured as CO₂, efflux werestudied at various soil water potentials, temperatures, androot ages for individual, attached roots of the barrel cactusFerocactus acanthodes and the platyopuntia Opuntia ficus-indica.The daily patterns of root respiration for both establishedroots and rain roots followed the daily patterns of root temperature.Root respiration increased when root temperature was raisedfrom 5 °C to 50 °C for F. acanthodes and from 5 °Cto 55 °C for O. ficus-indica; at 60 °C root respirationdecreased 50° from the maximum for F. acanthodes and decreased25° for O. ficus-indica. Root respiration per unit d. wtdecreased with root age for both species, especially for rainroots. Root respiration rates for rain roots were reduced tozero at a soil water potential (     

Chilling Damage to Photosynthesis in Young Zea mays: I. EFFECTS OF LIGHT AND TEMPERATURE VARIATION ON PHOTOSYNTHETIC CO2 ASSIMILATION

Carbon dioxide and water vapour exchanges of the second leafof Zea mays in controlled environment cuvettes were measuredin an open gas-exchange system, during and following subjectionto low temperature stress. Photosynthetic CO2 assimilation (Fc)decreased markedly with decrease in leaf temperature so thatFc at 5 °C was c. 7% of Fc at 20 °C. Fc continued todecline if leaf temperature was maintained at 5 °C, andwhen returned to 20 °C the leaf could not regain its previousFc. This chill-induced reduction in the capacity of the leafto assimilate CO2 was proportional to the duration of the chilland increased with water vapour pressure deficit and photonflux density (I_n). Six hours at 5 °C decreased Fc on returnto 20 °C, relative to Fc prior to treatment, by c. 10% indarkness and by c. 50% in a photon flux density approachingfull-sunlight (I_p = 1.5 mmol m^–2 s^–1). The degreeof reduction in Fc following chill treatment showed an almostlinear dependence on both the length and temperature of thechill. Chill treatments resulted in a decrease in both stomataland mesophyll conductances. Examination of the responses ofF_c to light and CO₂ concentration suggested that chill damageto the capacity for CO₂ assimilation resulted from effects onboth the light and CO₂ limited processes within photosynthesis. Key words: Chilling, Photosynthesis, Zea mays, Light-temperature interaction     

Effects of Temperature and Water Potential on Germination, Radicle Elongation and Emergence of Mungbean

Controlled environment experiments were performed to determinethe effects of temperature and water potential on germination,radicle elongation and emergence of mungbean (Vigna radiata(L.) Wilczek cv. IPB-M79-17-79). The effects of a range of constant temperatures (15–45°C) and water potentials (0 to –2.2 MPa) on germinationand radicle elongation rates were studied using an osmoticumtechnique, in which seeds were held against a semi-permeablemembrane sac containing a polyethylene glycol solution. Linearrelationships were established between median germination time(Gt₅₀) and water potential at different temperatures, and betweenreciprocal Gt₅₀ (germination rate) and temperature at differentwater potentials. Germination occurred at potentials as lowas –2.2 MPa at favourable temperatures (30–40 °C),but was fastest at 40 °C when water was not limiting, withan estimated base temperature (T_b) of about 10 °C. Subsequentradicle elongation, however, was restricted to a slightly narrowertemperature range and was fastest at 35 °C. The conceptof thermal time was used to develop an equation to model thecombined effects of water potential and temperature on germination.Predictions made using this model were compared with the actualgermination obtained in a related series of experiments in columnsof soil. Some differences observed suggested the additionalimportance of the seed/soil/water contact zone in influencingseed germination in soil. Seedling emergence appeared to reflectfurther the radicle elongation results by occurring within anarrower range of temperatures and water potentials than germination.Emergence had an estimated T_b of 12.6 °C and was fastestat 35 °C. A soil matric potential of not less than about–0.5 MPa at sowing was required to obtain 50% or moreseedling emergence. Key words: Germination, temperature, water potential     

H+ Efflux and Trans-Root Potential Measured while Increasing the Temperature of Solutions Bathing Excised Roots of Zea mays

Kennedy, C. D. and Gonsalves, F. A. N. 1988. H⁺ efflux and trans-rootpotential measured while increasing the temperature of solutionsbathing excised roots of Zea mays.—J. exp. Bot. 39: 37–49. Novel temperature-ramp procedures have been used to measureH⁺ efflux and trans-root potential of excised roots of Zea mays(var. Fronica). Two types of experiment were performed: (1),increasing temperature from 17°C, and (2), pre-cooling theroots to 1°C before starting the temperature ramp. The ratesof increase of temperature for H⁺ efflux and trans-root potentialexperiments were 0·5 and 2·1°C min^–1respectively The H⁺ scans revealed strong sharp maxima at 30°C and 32°C,for non-pre-cooled and pre-cooled roots respectively, the latterbeing significantly smaller. The trans-root potential scansfor the pre-cooled roots showed a corresponding maximum at 30°C,which was inhibited by KCN (1-0 mmol dm^–3) with or withoutSHAM (10 mmol dm^–3), or Hg²⁺ (1, 10, 100 µmol dm^–3)in the bathing solutions. Some of the evidence suggests thatthese maxima are associated with electrogenic H⁺ pumping, mediatedby a plasma membrane-bound ATPase. However, no correspondingmaximum was observed in the trans-root potential scans for non-pre-cooledroots, the potential remaining at about — 75 m V from20°C to 35°C. As there is a 7-fold increase in H⁺ effluxbetween 20°C and 30°C, the relationship between netH+ efflux and electrogenic proton pumping in these roots isby no means clear. Some possibilities are considered here. Pre-cooled and non-pre-cooled roots show clear maxima in thetrans-root potential scans at about 46°C, at which temperaturethere is a slight net H⁺ influx. This, and other less prominentfeatures observed, are briefly discussed. Key words: H⁺ efflux, trans-root potential, temperature-ramp procedure, Zea mays, roots     

Differences in the Interacting Effects of Light and Temperature on Growth of Four Species in the Vegetative Phase

A comparative study, employing the concepts of growth analysis,has been made of the varying responses in the early vegetativephase of Gossypium hirsutum, Helianthus annuus, Phaseolus vulgaris,and Zea mays to combinations of light intensity (1.08, 2.16,3.24, 4.32, and 5.4 x 10⁴ lx—photoperiod 14 h) and constantdiurnal air temperatures (10, 15, 20, 25, 30, and 35 °C).Depending on the combination of treatments, the temperatureof the internal tissues departed from air temperature by 6.9to 1.4 °C: so only the internal temperatures are cited here. For each species there are complex interactions between theeffects of light and temperature on the net assimilation rate,the leaf-area ratio, and the relative growth-rates of plantweight and leaf area. The magnitude of the changes induced bythe two factors vary both with the growth component and thespecies. The temperature responses are maximal up to 20–5°C while at the highest temperatures they may be negative.The temperature coefficients for leaf-area ratio are consistentlyless than those of the other three components: here betweenspecies the coefficients over 10–20 °C vary by a factorof 9.6, 5.4, and 5.1 for the rates of gain in plant weight andleaf area and the net assimilation rate, while the orderingwithin each growth component is species dependent. Under conditions of optimal temperature the relative growth-rateand net assimilation rate progressively increase, accordingto the species, up to either 4.32 or 5.4x 10⁴ lx. The leaf-arearatio is always largest at the lowest intensity. The level oflight at which the rate of gain in leaf area reaches a maximumranges from 2.16x 10⁴ lx for Phaseolus to between 4.32 and 5.40x10⁴ lx for Gossypium. The highest relative growth-rate and net assimilation rate ofHelianthus exceed those of Zea substantially. Indeed the maximalassimilation rate for Helianthus of 2.10 g dm^–2 week^–1is the highest ever recorded under field or controlled conditions.Possible reasons for this reversal of the photosynthetic potentialsof the two species observed by previous workers are discussed.     

The Effects of Root Cooling and the Light--Dark Transition on Growth of Primary Leaves of Phaseolus vulgaris L.

Use of Rotary Variable Displacement Transducers attached togrowing primary leaves of Phaseolus vulgaris has shown thatwhen the root systems were rapidly cooled from 23°C to 10°Cleaf extension rate fell to very low values within a few minutes.When the root systems were returned to 23°C leaf extensionincreased almost immediately to overshoot the control rateswithin 5–10 min, before declining to control values overthe next 50–60 min. When lights went off at the end ofthe day cycle there was an immediate and very large increasein leaf extension rate in both root-cooled and control plants;the rate then slowly declined over the next 60 min. This effectwas seen when the photoperiod was artificially shortened orlengthened and was reduced in magnitude when the photoperiodwas ended gradually by dimming the lights. The effect was notattributable to effects on leaf temperature but appears to bethe result of an endogenous rhythm interacting with the endof the photoperiod. At the beginning of the photoperiod therewas a gradual reduction in leaf extension rate occurring over30–45 min. Key words: Phaseolus vulgaris, leaf growth, extension rate, root cooling, wall extensibility, turgor     

The Influence of Temperature on Root Hair Infection of Trifolium parviflorum and T. glomeratum by Root Nodule Bacteria: I. THE EFFECTS OF CONSTANT ROOT TEMPERATURE ON INFECTION AND RELATED ASPECTS OF PLANT DEVELOPMENT

The onset and rate of infection in root hairs of T. parviflorumand T. glomeratum inoculated with Rhizobium trifolli strain5 varied much with root temperature. At moderate root temperature(18, 24, and 30 °C) infections were initiated earlier andin larger numbers than at low (6 and 12 °C) or moderatelyhigh (36 °C) temperatures. Both species showed a broad temperatureoptimum between 18 and 30 °C. The site of thread initiation(apically or laterally in a hair) was independent of temperature,as was also the proportion of successful threads penetratingthe root cortex, which increased with seedling age. Threadsgrew more slowly at low temperatures. The size of hair nucleinear infection threads remained unaffected by temperature, butnuclei associated with laterally originating threads were largerthan those associated with apical threads. Infection was non-randomly distributed along the main root atall temperatures. More zones of infection were found at moderatetemperatures than at temperature extremes (6–12 or 36°C). Probit plots of numbers of infections for individualplants were steplike, the linear sloping parts correspondingto normal distributions of infection within zones. Between 18 and 30 °C numbers of infections increased exponentiallyin two phases, the first more rapid phase ending at about thetime nodules appeared. A model devised for the infection processand fitted to the data suggested the existence of two kindsof infections: primary ones occuring randomly at a slow rateand probably not affected by temperature and secondary infectionsthat appeared to increase with rising temperatures in the range12 to 30 °C. Nodule numbers were relatively more sensitive to high and lowtemperatures than infection. The numbers of infections and nodulesand the root lengths of T. parviflorum were twice those of T.glomeratum except at the temperature extremes. Numbers of infectionswere otherwise unrelated to root length or cotyledon or leafareas. The development of lateral organs (primordia, lateralroots, and nodules) was reduced at temperatures below 18 °Cand above 30 °C.     

Depletion of Oxygen in Roots of Wheat and Maize by Respiration Following Encasement in Olive Oil

The extent to which isolated root systems of 14-d-old seedlingsof wheat (Triticum aestivum) and maize (Zea mays) were depletedof oxygen by respiration was measured after immersing them inoutgassed olive oil to exclude oxygen entry from the air. Atintervals over 45 min, gas from the roots was removed underpartial vacuum and oxygen partial pressures measured by gaschromatography. Contrary to earlier findings (Erdmann and Wiedenroth,1988), roots were able to utilize almost all of their oxygenwithin 20 min at 25°C, including that dissolved in the thinwater covering interposed between roots and oil. The rate ofaerobic respiration could be estimated readily from the timecourse of oxygen depletion.Copyright 1993, 1999 Academic Press Triticum aestivum, Zea mays, roots, anaerobiosis, respiration, oxygen, methods     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.). 1. Response of Leaf Extension to Temperature during the Transition from Vegetative to Reproductive Growth

The potential for leaf extension of plants of Lolium perennecv S24 growing in small artificial communities under naturalconditions was measured as the plants progressed from the vegetativeto the reproductive state In two consecutive years, 1975 and 1976, ‘simulated swards’were sown in autumn and overwintered in an open, unheated glasshouseIndividual swards from the batch sown in 1975 were brought into a growth room on two occasions in spring 1976 to measuretheir potential for leaf extension at a range of temperatures(5–20 °C) Individual swards from the batch sown inautumn 1976 were brought in to the growth room on 15 occasionsbetween November 1976 and May 1977 and their potential for leafextension was measured at a single temperature of 15 °CFrequent dissections were made in both years to describe changesin the developing apex. The potential for leaf extension at 15 °C decreased fromc 17 mm day^–1 in November to c 10 mm day^–1 in mid-winter.In January, the potential rapidly increased threefold to reach30mm day^–1 by mid February The increase began coincidentwith the earliest stages of floral initiation and was completedby the time of double-ridge formation Spring-grown vegetativeplants, however, showed potential rates of < 20 mm day^–1at 15 °C The results are discussed in relation to reproductive developmentand to changes in the carbohydrate strategy of the plants inearly spring Lolium perenne L perennial ryegrass, leaf extension, temperature response     

Leaf Extension in Zea mays: III. FIELD MEASUREMENTS OF LEAF EXTENSION IN RESPONSE TO TEMPERATURE AND LEAF WATER POTENTIAL

Leaf extension rates of young maize plants were measured inthe field. Large diurnal fluctuations of air temperature fromabout 5—35°C caused extension rates to vary from 0·4mm h^–1– 3·6 mm h^–1. When the temperatureof the shoot apical meristem was kept at 30–34°C,extension rates remained constant, despite diurnal fluctuationsof air temperature. Leaf water potentials () above –8 or –9 bars hadlittle apparent effect on extension rate. It was concluded that gradients within leaves could account for this.     

Fertility of Deep-frozen Maize (Zea mays L.) Pollen

On average, 50 per cent of maize pollen grains can be kept viableand almost 30 per cent remain fertile for up to a year whenthe water content on a fresh weight basis is reduced to about30 per cent of the original and the pollen is stored at –76or –196 °C. Over this period no significant differencewas found between storage at either temperature. Zea mays L., maize, pollen, fertility, viability     

Osmotic Adjustment and Stomatal Response to Water Deficits in Maize

A pot experiment was carried out using five maize {Zea maysL.) cultivars under three soil moisture levels (MPa 0 to –0.05,–0.3 to –0.9 and –1.2 to –1.5) to investigatethe effects of water deficits on osmotic adjustment and stomatalconductance. The degree of leaf rolling and the sugar and nutrientconcentrations in leaf cell sap were measured. Leaf water potential and osmotic potential decreased and stomatalconductance decreased with increasing water deficits. Stomatalconductance correlated positively with leaf water potentialand osmotic potential. Degree of leaf rolling was lower in cultivarswhich maintained higher turgor. Osmotic adjustment of 0.08 to0.43 MPa was found under the lowest soil moisture level in fivecultivars used. Sugar and K were the major osmotic substancesin the maize plant. Sugar, K and Mg concentrations increasedunder water deficit, and correlated negatively with a decreasein osmotic potential. Key words: Zea mays L., leaf water relations, leaf rolling, osmotic adjustment, stomatal conductance, water deficit     

The Responsiveness to Temperature of the Extension Rates of Leaves of Wheat Growing in the Field under Different Levels of Nitrogen Fertilizer

The response of the rates of extension (LER) of wheat leaves(Triticum aestivum cv. Gamenya) to temperatures maintained fora short period was measured by changing the temperature of theextension zone and recording the changes in leaf length. Therange of temperatures used was from 4-38 °C. The LER ofall leaves responded to increases in temperature as field temperatureswere suboptimal. The data obtained from several series of measurementsover different ranges of temperature were combined to producea general response curve. The minimum temperature for LER wasconsidered to be approximately 0 °C, the optimum was 28.4°C, while the maximum was in excess of 38 °C. The responsivenessof LER to temperature, measured by the Q₁₀, declined exponentiallyfrom >6 at 5 °C to 2 at 20 °C. The Q₁₀ at 15 °Cwas not affected by nitrogen supply.     

A Relationship between Membrane Permeability and Ethylene Production at High Temperature in Leaf Tissue of Phaseolus vulgaris L.

Leaf discs cut from primary leaves of Phaseolus vulgaris L cvMasterpiece were incubated at temperatures higher than the growthtemperature of 25 °C Both basal and wound ethylene productionincreased up to temperatures of 35–37 5 °C, thereafterdeclining rapidly There was no detectable ethylene productionat temperatures above 42 5 °C Exposure of leaf discs tohigh temperature for 60 mm resulted in a large production ofwound ethylene when they were returned to 25 °C The magnitudeof ethylene production was related to the initial incubationtemperature as was the length of the lag period before maximumproduction was achieved The results are discussed in relationto the requirement for continued membrane integrity for ethyleneproduction ethylene, temperature, membrane permeability, Phaseolus vulgaris L, dwarf bean