Studies on Extension Growth in Coleoptile Sections: II. The Effects of High Concentrations of {beta}-indolylacetic Acid on Section Growth

Using wheat coleoptile sections it has been shown that the treatmentgiven between cutting them from the coleoptile and placing themin the test solution greatly affects the early growth-rates. After growing sections in high concentrations of IAA for somehours it is necessary to give a considerable number of washingsto free them from surplus IAA; if these sections are then grownin water they reach lengths greater than those of comparablesections grown in water all the time, and in some cases greaterthan those attained in ‘optimum’ IAA concentrations. There is no suggestion in the experiments described that highconcentrations of IAA result in initial growth-rates higherthan those observed in ‘optimum’ concentrations,and at very high concentrations reduced early growth-rates areindicated. These results, and those described for lower IAA concentrationsin the first paper of this series, have some bearing on theapplication of the enzyme kinetic theory to auxin-induced growth,and this is considered in the discussion.     

The Effect of High Concentrations of Auxin on the Extension of Avena Coleoptile Sections

The response of Avena coleoptile sections to high concentrationsof auxin has been determined in the absence of all additivesexcept sucrose. In most experiments the growth-time curves with75 p.p.m. IAA showed two linear phases. In the first phase,which lasted for only 2–4 hours, extension was as rapidwith 75 p.p.m. IAA as with 5 p.p.m. IAA. This rapid initialexpansion phase was then succeeded by a second phase which persistedfor at least 20 hours. During this second linear phase the growth-ratewith 75 p.p.m. IAA was lower than with an auxin concentrationof 5 p.p.m. In some experiments the first phase was absent andonly the second phase was present. The response of sections to high concentrations of auxin wasnot influenced by the presence of buffers or absorbable cations.Omission of sucrose or the presence of moderate amounts of ethanolcaused the resulting growth curves to be non-linear. The rate of uptake of auxin into the tissues was dependent onthe auxin concentration and was constant for at least 24 hours.     

The Effects of Indole-3-Acetic Acid and 2:4-Dichlorophenoxyacetic Acid on the Growth Rate and Endogenous Rhythm of Intact AvenaColeoptiles

The rates of elongation of the coleoptiles of Avena seedlings,subjected to intermittent immersion in solutions of IAA or 2:4-Dfor various total periods, were determined from measurementsof photographs taken every hour by infra-red radiation. Immersion in 17·5 mg./l. IAA for 1–5 hours causeda large increase in the growth rate followed by a depression.When the seedlings were immersed in 8·75 mg./l. IAA forperiods of 12 or 24 hours the depression was partially overcomeso long as the treatment was continued. Absorption of additionalIAA by the coleoptiles reduced their geotropic sensitivity. Penetration of 2:4-D (sodium salt) into the coleoptiles wasslower than that of IAA and the resulting stimulation of thegrowth rate was less, particularly in unbuffered solutions.After the treatment the growth rate declined slowly to aboutthe normal value. Results with coleoptiles were very similar to those previouslyobtained with rhizomes of Aegopodium and suggest that inhibitionof growth following stimulation by IAA may be of general occurrence.Possible causes of the inhibition are discussed and a comparisonis made between the results with intact coleoptiles and observationsmade by others on coleoptile sections. Temporary immersion of the seedlings in auxin solutions depressedthe rate of elongation of the primary leaf while it increasedthat of the coleoptile. It caused little disturbance of theendogenous rhythm induced by change from light to darkness.The suggestion that such rhythms can be explained in terms ofvariation in concentration of IAA-oxidase is not supported.     

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     

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)     

Auxin Balance in the Avena Coleoptile

Coleoptiles of Avena possessed the capacity to degrade infiltrated indole-3-acetic acid (IAA). This activity decreased along the length of the coleoptile from apex to base on the bases of fresh weight, dry weight and protein; the apical 1 cm segment degraded more IAA than segments from other parts of the coleoptile. The naturally occurring inhibitor of the IAA oxidase activity increased in concentration up to 20 mm from the coleoptile apex; beyond, it decreased gradually towards the base. The spatial distribution of this inhibitor does not explain the gradient in IAA oxidase activity. Growth in length of the coleoptile and the IAA inactivating capacity of the apical 1 cm segment, increased 5- and 4,4-fold, respectively, between the ages of 70 and 130 h; but auxin secretion into agar platelets by the apical 2 mm of the coleoptile registered only a 2.7-fold increase. Deseeding and derooting the seedlings reduced the subsequent growth, diffusible auxin content and the IAA oxidase activity of the coleoptiles; derooting proved to be more deleterious than deseeding. A parallel reduction was evident in auxin content and IAA degrading activity following these treatments. Application of the cytokinin 6-benzylaminopurine (BAP) to coleoptiles of derooted seedlings failed to influence their capacity to degrade IAA. Nor was the activity of the aldehyde oxidase, which converts indole-3-acetaldehyde (IAAld) to IAA, affected by such treatment.     

Carbon Dioxide Effects on Auxin Responses of Coleoptile Sections

In the wheat cylinder bioassay technique as previously usedhere 5 sections have been enclosed in a 2 x 38 in, assay tubetogether with 0.5 ml. of the test solution. A method developedfor estimating the amount of carbon dioxide which accumulatesin these tubes through the respiration of the enclosed sectionshas shown that the level can rise to 20 per cent. after 24 hrs.at 25°C. In the presence of a 100 p.p.m. IAA(6x10^-4M.) testsolution, growth of 5 enclosed sections is depressed from 8hrs. onwardas and they eventually shrink, releasing their accumulatedIAA back into the solution. The growth of sections under various gas mixtures of carbondioxide in air has also been followed and these experimentsshow that section length is reduced approximately lineraly withrespect to increasing carbon dioxide concentration up to 20per cent. in air, both in the presence and absence of a 100p.p.m. IAA solution. The slope of the fitted regression line,however, is much steeper when the test solution contains IAA—i.e.there is a large interaction. In the presence of IAA, growth-time data show that a reductionin the growth rate, as compared with that in normal air, canbe detected after only 4 hrs, at the highest carbon dioxideconcentration. In the absence of IAA, high concentrations ofcarbon dioxide accelerate growth during the first 8 hrs. ofthe assay but depress it later. The mechanism of action of this interaction is unknown but itis not shown at very high concentrations of IAA, e.g. 1,000p.p.m. (6x10^-3M.).     

The Growth-time Relationships of the Auxin- induced Growth in Avena Coleoptile Sections

1. 1. The growth rate of Avena coleoptile sections in the presenceof indoleacetic acid (IAA) is constant with time over a widerange of time intervals and IAA concentrations.
2. 2. Constancyof growth rate is dependent upon the maintenanceof constantconditions in which the concentration of IAA availableto thesection remains the chief factor limiting growth rate.
3. 3.Control of the pH of the medium in which the sections aregrownis essential to the maintenance of constant growth rate,particularlyin the presence of high concentrations of IAA.
4. 4. The lagperiod in establishment of steady growth rate bysections inthe presence of IAA is less than 10 minutes andis not detectableby present methods of measurement.

     

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)     

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.     

Changes in Growth, Auxin- and Ribonucleic Acid Metabolism in Wheat Coleoptile Sections Following Pulse Treatment with Indole-3-Acetic Acid

Growth reactions of wbeat coleoptile sections following a brief pretreament in indole-3-acetic acid (LAA) were studied. The growth versus concentration curves 24 hours after the treatment showed a minimum value surrounded by bigber values. The minimum was never at concentrations lower than 10^-5M lAA but it could be found at higher concentrations after short pretreatment periods. The growth versus time curves reveated that the hormone treatment cansed the growth rate initially to increase but later on to decrease. The decrease was followed by a second increase for some treatments. Analysis of IAA content after the pretreatment showed that the attered growth patterns could be ascribed to declining auxin content with time, but not to thc actual concentration in the sections. The results indicate that the metabolic activation brought about by IAA leads to its own disappearance. Such a phenomenon was mirroretl in effects of IAA on hte net synthesis of ribonucleic acid.     

Suppression of (1→3),(1→4)-β-d-Glucan Turnover during Light-Induced Inhibition of Rice Coleoptile Growth

d -Glucan contents and (1→3),(1→4)-β-d-glucan hydrolase activity increased in the faster phase of coleoptile growth, then declined under both light and dark conditions. The relative glucan content in the cell wall showed a good correlation with the increment of coleoptile length. Strong correlations were also observed among the increment of coleoptile length, the decrease in the level of the glucans, and the relative activity of the glucanase in the cell wall of light- and dark-grown coleoptiles except for those values in the early stage of coleoptile growth, supporting a hypothesis that the turnover of the glucans is one of the important factors which regulate rice coleoptile growth. The levels of the glucans and the glucanase activity were always lower in the cell wall of coleoptiles grown in the light than those in darkness during the experimental period. These results suggest that light irradiation inhibits both the synthesis and the breakdown of the glucans, causing a decrease in the capacity of the cell wall to extend, thereby inducing growth suppression in rice coleoptiles. Received 24 September 1998/ Accepted in revised form 28 December 1998     

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.

     

Growth Regulation of Dark-grown Dwarf Barley Coleoptile by the Endogenous IAA Content

Growth curves of dark-grown coleoptiles of 11 isogenic coleoptilardwarf strains of barley (Hordeum vulagare L. cv. Akashinriki:uzu, 5, 77, 97, 105, 125, 131, 133, 136, 145 and 148) were simulatedwith a logistic equation and the endogenous IAA contents ofthe barley strains were determined. Growth analysis of the dwarfbarley coleoptiles revealed that the final coleoptile lengthwas correlated with the growth rate on the 2nd day after germination(r=0.897), when the growth rate was about maximum. The endogenousIAA Content of the barley strains, measured fluorometrically,indicated that on the 2nd day, the dwarf strains contained lessendogenous IAA than the normal Strain. The IAA content on the2nd day was correlated to the growth rate on the 2nd day (r=0.907,except for Strain 145) and the final coleoptile length (r=0.933,except for strains 77 and 145). The correlation, however, wasnot significant on the 3rd day. These results suggested thatthe dwarfism of the dark-grown coleoptiles of the barley Strainsexamined is primarily controlled by the endogenous IAA content. ¹ Present address: Department of Biology, Faculty of Science,Osaka City University, Osaka 558, Japan. (Received February 1, 1982; Accepted April 13, 1982)     

Morphological and Anatomical Characterization of the Coleoptile of Triticum aestivum with Regard to the Evolution of Forms with Different Ploidy Levels

Details of the morphology and anatomy of the coleoptile of wheatplants are given; these have not been described adequately previously.The investigations focussed on the hexaploid summer wheat Triticumaestivum L. cv. Hatri, and three taxa with different ploidylevels. In darkness the longitudinal growth of the coleoptile was delayedby nearly 24 h but the final length, reached after 120 h, wasdouble that of coleoptiles of plants cultivated under continuouslight. As soon as the coleoptile has grown, the primary leafpushes through a pore pre-formed during the meristematic stageand located 1–1·5 mm behind the apex. The poreis stabilized mechanically by anastomosing of the originallyfree ends of the vascular bundles, as well as by increased lignificationin this region. The species investigated differ in length, f.wt and d. wt, size of epidermal cells, and especially in thesize of guard cells of the coleoptiles. The number of parenchymalayers, however, shows no specificity. Triticum aestivum L. cv. Hatri, wheat, coleoptile, morphology, anatomy, Aegilops spp     

Studies on Extension Growth in Coleptile Sections: III. THE INTERACTION OF TEMPERATURE AND {beta}-INDOLYLACETIC ACID ON SECTION GROWTH

Wheat coleoptile sections were grown at various temperaturesfrom 1–40°C. in water and solutions of the sodiumsalt of ß-indolylacetic acid (NaIAA) ranging fromlow to very high concentrations (0.001 p.p.m. to 10,000 p.p.m.). Growth-time curves are sigmoid, obviously so under conditionsunfavourable to growth, less clearly as conditions either oftemperature or growth-substance supply are improved, the pointof inflection moving nearer to zero time. The growth-rate isreduced by temperatures below the optimum, and although growthcontinues for a longer time, final section length is also reduced.Such reductions are also brought about by sub-optimal levelsof NaIAA and by supra-optimal levels of both factors. In the test method employed, sections are grown in closed tubesof small volume, and after a time lose turgidity and becomewaterlogged; the onset of this condition is accelerated by hightemperature or growth-substance concentration, and greatly delayedby low temperature. Under comparable conditions, but with thetubes uncorked, flaccidity is delayed even at high temperatureor high concentration of NaIAA. If sections are transferred from one temperature to anotherthe change in growth-rate is abrupt; effects of transfer onfinal length are complex. A contour diagram represents the idealized relationships betweensection growth and temperature and NaIAA supply over their wholerange of effectiveness. Relations between temperature and growthin other systems are briefly discussed.     

The Extension Growth-Time Relationship for Avena Coleoptile Sections

1. The growth rates of coleoptile segments supplied with indole-3-aceticacid is not constant with time, but, when the IAA concentrationis high, decreases very rapidly.
2. With sufficiently high concentrationof IAA, the initial rapidgrowth may be eventually followedby shrinkage of the tissue.
3. The relation between initial rateof growth and auxin concentrationis not significantly differentfrom hyperbolic.
4. The significance of these facts in relationto kinetics of auxinaction is discussed.

     

Gravitropism and Phototropism of Maize Coleoptiles: Evaluation of the Cholodny-Went Theory Through Effects of Auxin Application and Decapitation

It was investigated whether or not gravitropism and phototropismof maize (Zea mays L.) coleoptiles behave as predicted by theCholodny-Went theory in response to auxin application, decapitationand combinations of these treatments. Gravitropism was inducedat an angle of 30° from the vertical, and phototropism,by a pulse of unilateral blue light. Either tropism of the coleoptilewas inhibited by IAA, applied as a ring of IAA-lanolin pasteto its sub-apical part, and by decapitation. The dose-responsecurves for the effects of applied IAA on tropisms and growthof intact coleoptiles as well as the time courses of tropismsinduced in decapitated coleoptiles could be explained by thethree conclusions in the literature: (1) the tip of the coleoptileis the site of auxin production, (2) lateral translocation ofauxin in gravitropism occurs along the length of the coleoptile,and (3) lateral translocation of auxin in phototropism occursin the coleoptile tip. By examining the effects of decapitationmade at different distances from the top and of IAA appliedto the cut surface of decapitated coleoptiles, it was indicatedthat auxin is produced in the apical 1 mm zone of an intactcoleoptile and that lateral auxin translocation for phototropismtakes place in an apical part that somewhat exceeds the zoneof auxin production. (Received October 14, 1994; Accepted December 26, 1994)     

Time-dependent Changes in the Auxin Sensitivity of Coleoptile Segments: Apparent Sensory Adaptation

When segments are excised from corn (Zea mays L.) coleoptiles they exhibit a very low rate of elongation for about 3.5 hours. A strong increase in growth rate (the spontaneous growth response) then occurs and persists for many hours. During the latent period preceding the spontaneous growth response there is an apparent increase with time in the sensitivity of the segments to indoleacetic acid (IAA). This increase in sensitivity is expressed as a 2- to 3-fold increase in the magnitude of the growth response to low levels of IAA and a 3-fold decrease in the latent period of the response during the first 3 hours following excision. A similar increase in sensitivity to low levels of IAA is noted if application of IAA is timed from the point of termination of a previous exposure to the hormone. Since the increase in responsiveness to low levels of IAA is not paralleled by an increase in the rate of uptake of the hormone, the data may be interpreted as evidence for a type of time-dependent sensory adaptation to auxin. The IAA dose-response relationship also changes with time, and there is indirect evidence that an auxin-dependent inhibitor may influence the expression of the apparent sensory adaptation to auxin.     

Inhibition of the Coleoptile Growth in Avena sativa by Endosperm Removal--Changes in Auxin, Osmotica and Cell Wall

Removal of the endosperm from 84-h-old etiolated oat seedlingsstrongly retarded the subsequent growth of coleoptiles. Thecontribution of the endosperm to coleoptile growth was studied.Endosperm removal was found to: (1) decrease the endogenouslevel of indole-3-acetic acid (IAA) in the coleoptile tip. IAAapplied to the coleoptile tip stimulated coleoptile growth inseedlings with and without the endosperm. The sensitivity ofthe coleoptile to a suboptimal concentration of IAA was higherin seedlings without the endosperm than in intact ones. At theoptimal concentration of IAA, however, the final length of thecoleoptile was larger in intact seedlings than in those withoutthe endosperm. (2) decrease the concentration of the solublesugars and amino acids in the cell sap. (3) retard the increasein the amount of polysaccharides in the cell wall of the coleoptile,particularly noncellulosic ones. (4) make the cell wall mechanicallyrigid according to stress-relaxation analysis of the cell wall.(5) induce an increase in the osmotic potential of the coleoptilecell sap. From these results, it was concluded that the endosperm suppliesthe coleoptile with IAA, sugars and amino acids, thus promotingcoleoptile growth. (Received September 24, 1987; Accepted February 3, 1988)