The Relationship between the Growth of the Primary Leaf and of the Coleoptile in Seedlings of Avena and Triticum

Partial inhibition of extension growth of the primary leaf occurswhen whole Triticum seedlings are immersed in aerated solutionsof IAA but is replaced by growth promotion when sucrose is addedto the external solution. In seedlings in which the coleoptilehas been excised, IAA increases the growth of the leaf bothwith and without additional sucrose. Inhibition of the leaf by moderate concentrations of IAA nolonger occurs when the seedling is detached from the endosperm.Sucrose added to the external solution raised the percentageelongation of the coleoptile almost to the level of that attainedin intact seedlings without additional carbohydrate. It alsoenabled the leaf to show a positive growth response with IAA. The results indicate that in intact seedlings treated with IAAthe growth of the primary leaf is markedly diminished owingto diversion of carbohydrate to the coleoptile if the growthof the latter is promoted as a result of the treatment. Whenthe competition of the coleoptile for carbohydrate is diminishedor eliminated, acceleration of the growth of the primary leafby IAA becomes apparent. In addition to the endogenous rhythm, with a period close to24 hours, induced in the growth-rate of the coleoptile whenseedlings of Avena are transferred from red light to darkness,a similar rhythm, with a slightly longer period, is inducedin the growth-rate of the primary leaf. This rhythm persistsin elongating leaves so long as they remain within the coleoptile.It can be recorded for at least 100 hours in deseeded seedlings. When intact seedlings of Avena are immersed for one hour inrelatively high concentrations of IAA and then transferred todistilled water for 18 hours, the elongation of the coleoptileis greater and the inhibition of the leaf is less than whenthey are transferred to humid air. Sections of the leaf of Triticum showed a slight increase inelongation in concentrations of IAA up to 5 mg./l., but no evidencewas obtained that sections of leaf and coleoptile exert any.influenceon each other's elongation when floated together on solutionsof IAA.     

The Effects of Externally Applied 3-indolylacetic Acid on Phototropic Induction and Response in the Coleoptile of Avena

The effects of intermittent immersion of Avena seedlings insolutions of IAA on the response of the coleoptiles to unilateralillumination in the region of that producing the second positivecurvature were studied by means of automatic time-lapse photographywhich enabled the growth-rate and curvature to be recorded simultaneously. Phototropic induction occurred even after the coleoptiles hadabsorbed sufficient IAA from a 10^-4 M. solution to raise theirrate of elongation to about twice the normal value. Phototropiccurvature, which had been temporarily inhibited by a curvaturein the opposite direction induced by the IAA, became evidentas soon as this curvature had ceased to operate. In coleoptiles, supplied with IAA after the commencement ofa phototropic curvature, the response was temporarily suppressed.It was resumed as soon as the effects of the exogenous IAA haddisappeared. The ability of the coleoptiles to produce a slight phototropicresponse persisted even when their growth-rate had been greatlyreduced by previous removal of the endosperm. Increasing thegrowth-rate by supplying the starved seedlings with IAA or sucrose,separately or together, failed to increase the response. Decapitation did not prevent phototropic induction, but delayedthe onset of the response. Application of IAA by intermittentimmersion in a 0.1 mg./l. solution, after the decapitated coleoptileshad been exposed to unilateral illumination, increased the rateof growth but reduced the response. The results suggest that in these experiments phototropic inductionwas not mediated by any direct action of light on the displacement,inactivation, or rate of synthesis of an endogenous auxin. Theyare in agreement with the hypothesis that the stimulus causedan asymmetrical distribution of a co-factor of auxin.     

Cooperation of Ethylene and Auxin in the Growth Regulation of Rice Coleoptile Segments

Elongation of coleoptile segments, having or not having a tip,excised from rice (Oryza sativa L. cv. Sasanishiki) seedlingswas promoted by exogenous ethylene above 0.3 µl l^–1as well as by IAA above 0.1 µM. Ethylene production ofdecapitated segments was stimulated by IAA above 1.0µM,and this was strongly inhibited by 1.0 µM AVG. AVG inhibitedthe IAA-stimulated elongation of the decapitated segment witha 4 h lag period, and this was completely recovered by ethyleneapplied at the concentration of 0.03 µl l^–1, whichhad no effect on elongation without exogenous IAA. The effectsof IAA and ethylene on elongation were additive. These factsshow that ethylene produced in response to IAA promotes ricecoleoptile elongation in concert with IAA, probably by prolongingthe possible duration of the IAA-stimulated elongation, butthat they act independently of each other. Moreover, AVG stronglyinhibited the endogenous growth of coleoptile segments withtips and this effect was nullified by the exogenous applicationof 0.03 µl l^–1 ethylene. These data imply that theelongation of intact rice coleoptiles may be regulated cooperativelyby endogenous ethylene and auxin in the same manner as foundin the IAA-stimulated elongation of the decapitated coleoptilesegments. Key words: oryza sativa, Ethylene, Auxin, Coleoptile growth     

Gravitropic Responses of Partially Decapitated Corn Coleoptiles with and without Applied [C]Indoleacetic Acid

The curvature of corn seedling (Zea mays L. Mo17 × B73) coleoptiles which had been half-decapitated and supplied with [¹⁴C]indoleacetic acid (IAA) (3.2 micromolar, 51 milliCuries per millimole) was determined during a 3-hour period of gravitational stimulation. Curvature of such half-decapitated coleoptiles was found to be similar in rate and extent to that of intact coleoptiles responding to gravity. Gravitational stimulation was accomplished by reorienting seedlings to a horizontal position, either up or down with respect to the removed half of the coleoptile tips.

The first set of experiments involved placing aluminum foil barriers along one of the two cut surfaces to restrict the movement of IAA into tissues. The initiation and extent of curvature of these half-decapitated coleoptiles was dependent upon the orientation of the removed half-tip and the accompanying barrier. The distribution of radioactivity from [¹⁴C] IAA after 3 hours indicated that the specific lateral movement of label was also dependent upon orientation of the removed half-tip of the coleoptile. A specific movement to the lower side of approximately 14% of the total recovered radioactivity was found in coleoptiles in which the [¹⁴C]IAA was supplied across a transverse cut surface. In contrast, specific movement of only 4% was found for application across a longitudinal cut surface.

A second series of experiments was conducted using 1.0 and 3.2 micromolar [¹⁴C]IAA (51 milliCuries per millimole) supplied to half-decapitated coleoptiles without inserted barriers. The 3.2 micromolar concentration adequately replaced the removed coleoptile half-tips in terms of straight growth, but it did not result in as much curvature as shown by coleoptiles of intact seedlings. The 1 micromolar concentration was not adequate to replace the removed half-tip in straight growth, but resulted in gravitropic curvature nearly as great as that produced by the higher concentration.

The data presented here suggest that strong auxin gradients are not produced in response to gravity stimulation based on the recovered radioactivity from [¹⁴C]IAA. However, it is evident that auxin is required for the development of normal gravitropic responses. It is possible, therefore, that an important early role of this movement is not to cause a large stimulation of growth on the lower side but to decrease growth on the upper side of a gravitropically responding coleoptile.

     

Distribution of Endogenous Indole-3-acetic Acid and Growth Inhibitor(s) in Phototropically Responding Oat Coleoptiles

The relationship between the flank growth of oat (Avena sativaL. cv. Victory) coleoptiles and the distribution of endogenousindole-3-acetic acid (IAA) and growth inhibitor(s) in the coleoptileswas studied for the second positive phototropic curvature inducedby a continuous unilateral illumination with white light (0.1W.m^–2). The phototropic curvature was caused by growthinhibition at the lighted side and growth promotion at the shadedside. Using electron capture detection gas chromatography, weanalyzed the distribution of endogenous IAA in phototropicallyresponding oat coleoptiles and found that the IAA was evenlydistributed over the lighted and shaded sides during the phototropicresponse; there was also no detectable difference in the amountsof IAA between phototropically stimulated and non-irradiatedcoleoptiles. By contrast, oat coleoptile straight-growth testresults showed that the amount of unknown acidic growth inhibitor(s),different from abscisic acid, increased in the lighted halfof the coleoptiles and decreased in the shaded half, as comparedto the amount in the non-irradiated half. These data suggestthat the phototropic curvature of oat coleoptile is inducedby a difference in lateral flank growth through a lateral gradientof endogenous growth inhibitor(s) rather than of IAA. (Received February 10, 1988; Accepted July 29, 1988)     

Inhibition of IAA-Induced Growth of Wheat Coleoptile Fragments by Chitin-Chitosan Oligomers

We studied the comparative effects of the phytohormone indolylacetic acid (IAA) and chitooligosaccharides (5–10 kDa, degree of deacetylation 35%) on the growth of coleoptile fragments of 3-day wheat germlings. IAA (10mg/l) stimulated elongation of coleoptile fragments by 80%, as compared to the control (water). Chitooligosaccharides at 0.01–1500 mg/l or higher did not exert auxin-like effects, but at 100 mg/l or higher, suppressed elongation of the coleoptile fragments, as compared to the control (water). Incubation of coleoptile fragments in solutions containing both IAA and chitooligosaccharides (100 mg/l or higher) suppressed their IAA-induced elongation, which correlated with the inhibition of growth of 3-day wheat germlings after wetting seeds in solutions of chitooligosaccharides. It has been proposed that the phytohormone-like properties of chitooligosaccharides can be related to changes in the endogenous balance of phytohormones in plants.     

Metabolism of [14C]Indole-3-Acetic Acid by Coleoptiles of Zea mays L.

Reverse-phase high-performance liquid chromatography was usedto analyse [¹⁴C]-labelled metabolites of indole-3-acetic acid(IAA) in coleoptile segments of Zeo mays seedlings. After incubationfor 2 h in 10^–2 mol m^–3 [2-¹⁴C]IAA, methanolic extractsof coleoptiles contained between six and ten radioactive compounds,one of which co-chromatographed with IAA. The metabolic productsin coleoptile extracts appeared to be similar to those in rootextracts, with an oxindole-3-acetic-acid-like component as theprincipal metabolite, but the rate of metabolism was slowerin coleoptile than in root segments. Decarboxylation did notappear to play a major role in the metabolism of exogenous IAAduring the short incubation periods. Moreover, external IAAconcentration had little effect on the pattern of metabolism.Coleoptile segments were also supplied with [¹⁴C]IAA from agardonor blocks placed at the apical ends, and agar receiver blockswere placed at the basal ends. After incubation for 4 h, theidentity of the single radioactive compound in the receiverblocks was shown to be IAA by both reverse-phase high-performanceliquid chromatography and gas chromatography-mass spectrometrytechniques. Key words: Zea mays, Coleoptile, High-performance liquid chromatography, Indole-3-acetic acid     

Stress relaxation properties of the cell wall of tissue segments under different growth conditions

Stress-relaxation parameters were compared under different experimentalconditions using 5th internode segments of light-grown pea seedlingsand coleoptile segments of dark-grown Avena seedlings. The followingresults were obtained. 1. In a short incubation period at 25?C, IAA caused a decreasein the minimum relaxation time, To, of the epidermal cell wallof pea internodes when it induced elongation; the optimum concentrationof IAA for decreasing To was 10 mg/liter. 2. At all concentrations of IAA used, 0.1–1000 mg/liter,the relationship between the To value of the epidermal cellwall peeled from segments incubated for 2 hr and the subsequentelongation rate in 2–3 hr incubation was linear, indicatingthat the To value of the cell wall at a certain time regulatesthe rate of the following elongation. 3. When segments of pea epicotyls or Avena coleoptiles wereincubated in mannitol solution of various concentrations inthe presence and absence of IAA and then allowed to grow inthe absence of both mannitol and IAA, the segments extendeddifferently depending upon the mannitol concentration, whichwas less than 0.3 M, given during preincubation. 4. The T_o and b (relaxation rate, S/log t) values were smallerin the cell wall of segments which extended more, than in thosewhich extended less. In this case, 0.2 M mannitol solution wasmost effective, since it inhibited IAA-induced elongation duringpre-incubation and the segments thus incubated extended themost afterward. 5. Extensibility, mm/gr, seemed to parallel the elongation whichhad occurred during pre-incubation, indicating that this value,contrary to To, represented at least partly the result of elongation. From these results we concluded that the growth rate to followis regulated by the minimum stress relaxation time, To, andpossibly by the relaxation rate, b, of the cell wall beforeextension, and these parameters may represent certain biochemicalmodifications of the cell wall components needed for cell extension. (Received August 12, 1974; )     

Effects of γ-irradiation on elongation and indole-3-acetic acid level of maize (Zea mays) coleoptiles

The effects of γ-irradiation on elongation and the level of indole-3-acetic acid (IAA) of maize (Zea mays) coleoptiles were investigated. When 3-day-old seedlings of maize were exposed to γ-radiation lower than 1 kGy, a temporal retardation of coleoptile elongation was induced. This retardation was at least partly ascribed to a temporal decrease in the amount of free IAA in coleoptile tips on the basis of the following facts: (1) the reactivity to IAA of the elongating coleoptile cells was not altered by irradiation; (2) endogenous IAA level in the tip of irradiated coleoptiles was at first unchanged, but then declined before returning to nearly the same level as that of the non-irradiated control; and (3) the amount of IAA that diffused from coleoptile tip sections showed a similar pattern to that of endogenous IAA. The rate of conversion between free and conjugated IAA was not significantly affected by irradiation. These results suggest that a temporal inhibition of maize coleoptile elongation induced by γ-irradiation can be ascribed to the reduction of endogenous IAA level in the coleoptile tip, and this may originate from the modulation in the rate of IAA biosynthesis or catabolism.     

The Effect of Oxygen and Turbulence on Elongation of Coleoptiles of Submergence-Tolerant and -Intolerant Rice Cultivars

Coleoptiles of rice (Oryza saliva L. ) elongated more rapidlyin stagnant solution than in water-saturated air: elongationrates in aerated solution were intermediate. Elongation wasstimulated between 170% to 390% in stagnant solution in 17 cultivarsscreened. On this basis, a relatively submergence-intolerantcultivar and two submergence-tolerant cultivars were studiedfurther. When seedlings were grown in a range of oxygen concentrationsimposed by bubbling solutions rapidly with gas, germinationand early coleoptile growth (up to 5.0 mm) were inhibited atlow oxygen concentrations, particularly in the submergence-intolerantcultivar. Weight per unit length declined with time in thiscultivar, even in mild oxygen deficits. However, later stagesof coleoptile elongation were unaffected by low oxygen supply:elongation proceeded at control rates even in anoxia. It isconcluded that processes unique to germination and early stagesof coleoptile elongation (possibly cell division) have greateroxygen requirements than cell extension. Rates of elongation in stagnant solution were much faster thanthose in any other treatment, including bubbled solutions atsimilar oxygen concentrations (0.125 mmol O₂ I_–1). Therefore,elongation appears to be stimulated by accumulation of endogenouscompounds which are dispersed in bubbled solutions and in air.The principal volatile compound was almost certainly ethylene,since Ag+ (an antagonist of ethylene) reduced the rate of elongationin stagnant solution by 53%. Differences in turgor pressure were only partly responsiblefor rapid coleoptile elongation observed in stagnant solution:there must have also been differences in cell wall propertiesbetween coleoptiles grown in stagnant and bubbled solution.The rapid elongation in stagnant solution hastens coleoptileemergence from the solution, thus establishing an adequate oxygensupply for the seedling. Leaf and root growth then proceeds.     

Short-Term Stimulation of Growth Induced by the Apical Application of IAA to Intact Maize Coleoptiles

Indole-3-acetic acid (IAA), mixed with lanolin, was appliedto the surface of defined parts of the intact coleoptile ofmaize (Zea mays L.) seedlings, and the elongation growth ofthe coleoptile zone was monitored for a period of 8 h. WhenIAA was applied to a subapical region, the growth of zones locatedbelow was stimulated only temporarily, although stimulationwas continuous when IAA was applied directly to the zone thatwas monitored for growth. This short-term stimulation was observedat all concentrations tested (0.03–30 mg of IAA per gof lanolin). The duration of stimulated growth, which variedfrom 1 to 5 h, was longer as the distance from the site of IAAapplication was reduced and the concentration of IAA was increased.Growth during the post-stimulation phase after application ofa high dose of IAA could not be enhanced again by applying IAAnear the site of the first application, but it was enhancedwhen IAA was applied directly to the zone used to measure growth.These results suggest that the supply of IAA from the site ofapplication to lower zones is controlled internally such thatthe elevated concentration of IAA in these zones returns tothe concentration before IAA application. Either the feedbackregulation of auxin transport or the feedforward regulationof auxin metabolism may account for the suggested control ofIAA supply. (Received May 18, 1995; Accepted October 20, 1995)     

Inhibitory action of red light on the growth of the maize mesocotyl: evaluation of the auxin hypothesis

Brief irradiation of 3-d-old maize (Zea mays L.) seedlings with red light (R; 180 J m^-2) inhibits elongation of the mesocotyl (70–80% inhibition in 8 h) and reduces its indole-3-acetic acid (IAA) content. The reduction in IAA content, apparent within a few hours, is the result of a reduction in the supply of IAA from the coleoptile unit (which includes the shoot apex and primary leaves). The fluence-response relationship for the inhibition of mesocotyl growth by R and far-red light closely resemble those for the reduction of the IAA supply from the coleoptile. The relationship between the concentration of IAA (1–10 M) supplied to the cut surface of the mesocotyl of seedlings with their coleoptile removed and the growth increment of the mesocotyl, measured after 4 h, is linear. The hypothesis that R inhibits mesocotyl growth mainly by reducing the IAA supply from the coleoptile is supported. However, mesocotyl growth in seedlings from which the coleoptiles have been removed is also inhibited by R (about 25% inhibition in 8 h). This inhibition is not related to changes in the IAA level, and not relieved by applied IAA. In intact seedlings, this effect may also participate in the inhibition of mesocotyl growth by R. Inhibition of cell division by R, whose mechanism is not known, will also result in reduced mesocotyl elongation especially in the long term (e.g. 24 h).Abbreviations FR far-red light - IAA indole-3-acetic acid - P_fr phytochrome in the far-red-absorbing form - P_r phytochrome in the red-absorbing form - R red light     

Relationship between Coleoptile Elongation and Alcoholic Fermentation in Rice Exposed to Anoxia. I. Importance of Treatment Conditions and Different Tissues

The relationship between coleoptile elongation and alcoholicfermentation of rice under anoxia is examined using seeds either:(a) N₂ flushed during submergence, (b) incubated in stagnantdeoxygenated agar at 0·1% w/v to simulate the stagnantconditions of waterlogged soil, or (c) incubated in waterloggedsoil. Coleoptile elongation growth was greater for N₂ flushing> stagnant agar > soil; seed survival was also greatestin this order over 1-5 d. Ethanol concentrations in coleoptiles and intact seeds (cv.IR42) were approximately 300 and 100 mol m^-3 respectively whenseeds were grown 3 d in stagnant agar, however 92% of the ethanolin seeds diffused into the external medium when solutions weremixed for 5-10 s. Coleoptile growth under anoxia was relatedto rates of ethanol synthesis (R_E) in different treatments;there was greater coleoptile growth and R_E for seeds in N₂ flushedsolutions than in stagnant deoxygenated agar. Coleoptile growthof individual seeds was also related to the R_E of each seedat 2-3 d after anoxia (r² = 0·46). Analysis of different tissues was important in evaluating growthand metabolism of coleoptiles. Although the coleoptile onlyaccounted for 0·7% of seed dry weight at 3 d after anoxia,it contained 21% of the ethanol produced by rice seeds. Therewere also three-fold higher rates of R_E on a fresh weight basisin expanding tissues in the base of the coleoptile relativeto the elongated tissues at the apex. Results are discussedin terms of the importance of environmental conditions usedto impose anoxia, quantification of R_E in specific tissues andthe possibility that under stagnant conditions high ethanolconcentrations in tissues may limit R_E and coleoptile growth.Copyright1994, 1999 Academic Press Anoxia, ethanol, alcoholic fermentation, Oryza sativa L., rice, submergence     

Growth Response of Triticum Coleoptiles to Non-dialysable Milk Fractions

Dialysed skimmed cows' milk was shown to promote the elongationof Triticum vulgare var. ‘Yogo’ coleoptiles. Theoptimum concentration of skimmed milk was found to be one partto 200 parts distilled water. Milk dialysate was shown to inhibitelongation but after the cations were removed to have no influenceupon elongation. Gelatin, ß-lactalbumin, phosphovitin, and glycoproteinfraction IV each present at a concentration of 30 mg./100 ml.,increased coleoptile elongation about 20 per cent. when comparedto a distilled-water control. Proteose-peptone and -lactalbumincaused a slight inhibition in elongation. These results are discussed in terms of possible factors suchas bacterial contamination or adsorbed growth-promoting substancesas agents causing these effects.     

The Role of the Coleoptile Apex in Controlling Organ Elongation: I. THE EFFECTS OF DECAPITATION AND APICAL INCISIONS

It has been widely accepted for over 50 years that the elongationrate of a coleoptile is dependent on the supply of auxin fromthe apex. The original coleoptile decapitation experiments whichprovided support for this view have been repeated but the measurementsof coleoptile elongation were made with greater temporal andspatial precision. The experiments confirm that Avena and Zeacoleoptile elongation is retarded by decapitation but the locationand timing of the growth rate changes are not consistent withthe hypothesis that decapitation reduces growth rate solelyby removing the major supply of auxin. Evidence is presentedthat wounding is the prime cause of the effects of decapitation.Data are also presented showing that the recovery of growthrate of coleoptiles after decapitation or wounding is not dearlyassociated with any events near the cut surface and hence thetraditional explanation of this phenomenon (‘regenerationof the physiological tip’) is misleading. Key words: Coleoptile, decapitation, apex     

The Effects of Certain Growth-regulating Substances on the Rhizomes of Aegopodium podagraria

1. The effects on elongation and geotropic behaviour of theimmersion of rhizomes of Aegopodium podagraria for periods ofminutes in solutions of ß-indolylacetic acid,2:4-dichiorophenoxyaceticacid, ascorbic acid, 2:3:5 -triiodobenzoic acid, and 2:4-dichloranisolewere investigated. 2. The experimental treatment was carried out in absence ofvisible light, the rhizomes being photographed every hour byinfra-red radiation. 3. After immersion in 10^– M. IAA, the rate of growth isapproximately doubled. About 5 hours later it declines to one-thirdor less of its original value and then gradually recovers. Thedepression in the growth rate can be reversed by a second immersionin IAA, indicating that it is not due to exhaustion of someother substance essential for growth. 4. Immersion in 10^–4 M. 2:4-D also stimulates growth,which then slowly returns to about the normal rate without showingany marked depression. 5. The possibility is suggested that IAA, which first accelerateselongation, is later converted into, or causes the productionof, an inhibitor. When successive immersions in IAA are repeatedevery 2 hours, the inhibition can be partially overcome so longas the treatment is continued. 6. Ascorbic acid slightly depresses the rate of elongation,but triiodobenzoic acid and dichloranisole produce no significanteffect. 7. Immersion of the horizontal rhizomes in IAA or 2:4-D causesthem to turn up. This suggests that an equilibrium is disturbedand supports the hypothesis that the diageotropic behaviourof these rhizomes is the result of a balance between the effectsof two opposing hormones.     

Epidermal Growth Factor Interacts with Indole-3-Acetic Acid and Promotes Coleoptile Growth

We used coleoptile sections of Avena sativa, Sorghum bicolor,and Zea mays seedlings to examine interactions between epidermalgrowth factor (EGF) and indole-3-acetic acid (IAA) that mayaffect plant growth and development. Our 24-h bioassays employedthree controls ranging in dilution from 10^–4 to 10^–8g ml^–1: (1) 50 mM potassium-phosphate buffer solution(pH=6.0), (2) bovine serum albumin, a nonspecific protein; and(3) IAA; plus two treatments: (1) mouse epidermal growth factor(EGF) ranging from 10^–6 to 10^–10gml^–1, and(2) EGF + IAA. In all three species growth in IAA, EGF, andEGF + IAA treatments showed significant increases over controls;EGF+IAA showed significant increases in growth over IAA alone.As the concentrations of IAA decreased, the EGF and IAA interactionbecame more pronounced. At the highest IAA concentrations, EGF+ IAA increased growth rates ca. 2% to 39%, whereas at lowerIAA concentrations EGF + IAA promoted growth as much as 121%,thereby lowering the normal IAA physiological set point up tothree or four orders of magnitude. Our data suggest that aninteraction between EGF and IAA may allow plants to recognizeand respond to animal biochemical messengers, resulting in changesin plant cell elongation that ultimately may alter plant growthpatterns. (Received April 27, 1994; Accepted September 5, 1994)     

Osmoregulation in Rice Coleoptile Elongation as Promoted by Cooperation between IAA and Ethylene

IAA-induced elongation of rice (Oryza sativa L. cv. Sasanishiki)coleoptiles is regulated by cooperation between IAA and ethyleneproduced in response to IAA. However, the presence of some solutes,such as K^$, Na^$, Rb^$, glucose and sucrose, in the incubationmedia was found to be indispensable for this cooperation. Withoutthose solutes, the IAA-induced elongation was not sustainedover a long time period. IAA caused increases in both the osmoticpotentials of the coleoptile cells and the extensibility oftheir cell wall. In epidermal cells of IAA-treated coleoptiles,the osmotic potential increased from –0.87 to –0.62MPa during a 4-h incubation with 1 mM KCl. Moreover, IAA promotedthe uptake of K^$ or Na^$ from the media into the coleoptiles.However, these effects of IAA were partially prevented by aminoethoxyvinylglycine(AVG), and all the AVG effects were completely nullified byethylene applied simultaneously and exogenously. Both IAA andethylene did not affect the wall yield stress. These resultssuggested that the long-term elongation induced by IAA in ricecoleoptile segments results from inhibiting increases in osmoticpotentials of their cells. The maintenance by IAA of low osmoticpotentials may be partly due to the promotive action of ethyleneproduced in response to IAA on the solute uptake from the media. (Received July 6, 1983; Accepted February 15, 1984)     

Ethylene and the Responses to Light of Rice Seedlings

The growth of rice seedlings (Oryza satira L.) in the presence of ethylene caused a change in the response to light of coleoptile elongation. In plants grown in air without added ethylene coleoptile elongation was promoted by red, far-red and yellow-green light only in very young seedlings; in older plants irradiation inhibited the growth of the coleoptile. The effect of growing plants in the presence of ethylene was to prolong the period during which light promoted coleoptile growth. Elongation of the first internode was inhibited by light whether or not the seedlings were grown in the presence of ethylene. A correlation existed between the growth effect of an irradiation and the initial decay rate of phytochrome which was established by the treatment. Regardless of wave length, light sources whose intensities were adjusted to produce a decay rate of about 10% per hour or less induced a moderate rate of coleoptile elongation which persisted for a relatively long period. Irradiation with red or yellow-green light of higher intensity which produced a higher rate of phytochrome decay induced a higher rate of coleoptile elongation, but growth stopped after several hours. Other observations, however, showed that one cannot establish a general simple correlation between the rate of elongation of rice coleoptiles under light and the status of measurable phytochrome in the plant.     

Auxin and Ethylene Control of Growth in Seedlings of Zea mays L. and Avena sativa L

The effects of applied ethylene on the growth of coleoptilesand mesocotyls of etiolated monocot seedlings (oat and maize)have been compared with those on the epicotyl of a dicot seedling(the etiolated pea). Significant inhibition of elongation by ethylene (10 µll^–1for 24 h) was found in intact seedlings of all three species,but lateral expansion growth was observed only in the pea internodeand oat mesocotyl tissue. The sensitivity of the growth of seedlingparts to ethylene is in the decreasing order pea internode,oat coleoptile and oat mesocotyl, with maize exhibiting theleast growth response. Although excised segments of mesocotyland coleoptile or pea internode all exhibit enhanced elongationgrowth in IAA solutions (10^–6–2 ? 10^–5 moll^–1), no consistent effects were found in ethylene. Ethyleneproduction in segments was significantly enhanced by applicationof auxin (IAA, 10^–5 mol l^–6 or less) in all tissuesexcept those of the eat mesocotyl. Segments of maize show a slow rate of metabolism of applied[2-¹⁴C]IAA (30 per cent converted to other metabolites within9 h) and a high capacity for polar auxin transport. Ethylene(10 µl l^–1 for 24 h) has little effect on eitherof these processes. The oat has a smaller capacity for polartransport than maize and the rate ef metabolism of auxin isas fast as in the pea (90 per cent metabolized in 6 h). Althoughethylene pretreatment does not change the rate of auxin metabolismin oat, there is a marked reduction in auxin transport. It is proposed that the insensitivity of maize seedlings toethylene is related to the supply and persistence of auxin whichcould protect the seedling against the effects of applied orendogenously produced ethylene. Although the mesocotyl of oatis sensitive to applied ethylene it may be in part protectedagainst ethylene in vivo by the absence of an auxin-enhancedethylene production system. The results are discussed in relationto a model for the auxin and ethylene control of cell growthin the pea.