Studies on the Geotropism of Roots:II.THE EFFECTS OF THE AUXIN ANTAGONIST {alpha}-(I-NAPHTHYLMETHYLSULPHIDE)PROPIONIC ACID (NMSP) AND ITS INTERACTIONS WITH APPLIED AUXINS

Previous experiments on the effects of auxins on the geotropicresponses of seedling pea roots (Audus and Brownbridge, 1957)have been extended using the ‘anti-auxin’ -(I-naphthylmethylsulphide)propionicacid (NMSP) alone and in combination with indole-3-acetic acid(IAA) and 2:4-dichlorophenoxyacetic acid (2:4-D). NMSP action differs from that of the auxins in that it reducesthe rate of curvature progressively as the concentration isincreased, irrespective of whether the overall extension growthof the roots is being stimulated (10 and 30 p.p.m.) or inhibited(100 p.p.m.). Correspondingly the reaction time is lengthenedby 25–50 per cent. in all concentrations. Studies of responsesin mixtures of growth-stimulating concentrations of NMSP (30p.p.m.) and growth-inhibiting concentrations of IAA (10^–8)and 2:4-D (3 x 10^–8) show that auxins and ‘antiauxins’are mutually antagonistic in most, if not all, their actionson growth and curvature. The results suggest that the anti-auxin NMSP may stimulate rootgrowth and inhibit curvature by interfering with the synthesisor distribution of a natural endogenous inhibitor, which isnot IAA. NMSP inhibition of root growth in high concentrationsmust, however, be exerted independently of this natural inhibitor.The mutual antagonisms shown between the auxins and NMSP arebest explained in terms of an interference with access to thegrowth centres; competitive action at the growth centres themselvesseems not to be involved.     

Factors Affecting the Abscission of Reproductive Organs in Yellow Lupins (Lupinus luteus L.): III. ENDOGENOUS GROWTH SUBSTANCES IN VIRUS-INFECTED AND HEALTHY PLANTS AND THEIR EFFECT ON ABSCISSION

Extracts of small and mature-size lupin pods yielded four substancesaffecting the growth of wheat-coleoptile sections: one acidpromotor (A), two acid inhibitors(B and X), and one neutralinhibitor(Y). Inhibitor B was extremely active, however, coleoptile sectionsshowed no signs of toxic effects; they resumed growth at a rapidrate after rinsing them and adding ß-indolylaceticand (IAA) to the medium. 1 µg of IAA was required to counteractthe effect of ‘B’ extracted from 230 mg. Of tissue.On an equal fresh weight basis the inhibiting action of ‘B’in lupin pods was 500–1,500 times more potent than thatof ‘inhibitor ß’ in etiolated pea seedlings. Small pods of plants infected with pea-mosaic virus yielded3 times the amount of ‘A’ of healthy plants (equivalentto 1 µg. IAA 0.3 µg. IAA per 25 g. of tissue respectively),and approximately the amount of ‘B’. Mature podsof virus-infected plants again yielded more‘A’,but also 2? times more ‘B’ than pods of healthyplants. Healthy pods yielded more ‘A’ than virus-infectedpods, and there was no difference in ‘X’. A lupin abscission test was developed and the effects of proximaland distal application of -naphthyl acetic acid (NAA) are presented,and discussed with respect to results of other abscission tests. ‘A’ accelerated abscission when applied proximally,and delayed or prevented it when applied distally. ‘B’strongly accelerated abscission when applied in either way.A possible mechanism explaining the abscission-inducing effectof developing pods on later flowers is discussed in terms ofthe substances ‘A’ and ‘B’. The partlyprevented abscission observed on virus-infected plants was foundto agree well with the proposed mechanism.     

Some Growth Regulators of Tobacco in Relation to the Symptoms of the Physiological Disease 'Frenching'

Extracrts of the shoot tips of normal and ‘frenched’tobacco plants were chemically separated into acidic, neutral,and basic ether–soluble fractions. On chromatograms ofthese, some plant growth regulators were assayed using the Avenacoleoptile section extension test. The acidic auxins and an acids and a neutral growth inhibitorwere found. One auxin, with the samew R_F value as indole-3-aceticacid, was four times more concentrated on normal as in ‘frenched’plants. No differences could be established between the twotypes of plants in regard to other growth regulators detected. It is argued that the symptoms of the physiological disease‘frenching’ could be explained in terms of a auxindeficiency.     

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     

Studies on the Geotropism of Roots: III. EFFECTS OF GEOTROPIC STIMULATION ON GROWTH-SUBSTANCE CONCENTRATIONS IN VICIA FABA ROOT TIPS

The auxins contained in 5-mm. tips of horizontal Vicia fabaroots have been compared with those in tips of vertical rootsafter cold ethanol extraction, paper-chromatographic separation,and Avena mesocotyl bioassay. At about the time curvature commencesin horizontal roots there is a marked increase in the contentof an auxin corresponding to ‘AP(ii)’ of pea roots(Rf 0.35–0.65 in isobutanol/methanol/water). There areindications that this is not due to its release from an inactivebound state but that it is either synthesized de novo or maybe converted from another auxin corresponding to ‘AP(iii)’of pea roots (Rf 0.75–1.0). The literature dealing with the auxins of geotropically stimulatedorgans is reassessed and it is concluded that, with the exceptionof the Avena coleoptile, there is very little evidence favouringa simple transport redistribution of auxin under gravity; themajority of the data favour an effect of gravity on auxin metabolism.     

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.     

Metabolism of Some Indole Auxins in Excised Tomato Roots

Indolylacetylaspartic acid (IAAsp) and possibly indolylacetylglutamicacid (IAG) are formed by exposure of excised tomato roots toIAA. Little ‘free’ IAA accumulates in the tissue.An unidentified substance reacting pink with nitricnitrite reagentis also formed. These substances are metabolized when IAA-treatedroots are transferred to auxin-free medium. IAAsp and IAA aresimilarly inhibitory to the growth of excised tomato roots.Excised tomato roots do not interconvert IAA and IAN. IAN-feedingleads to IAN accumulation and the appearance of indolylcarboxylicacid (ICA); transference to auxin-free medium causes a declinein the IAN activity but the ICA spot persists. The inhibitoryactivity of IAN is not due to its conversion to ICA. Excisedtomato and wheat roots respond very differently to externall-tryptophane but in neither case is there evidence of the conversionof tryptophane to ethyl acetate-soluble auxins.     

The Geoelectric Effect in Plant Shoots: III. DEPENDENCE UPON AUXIN CONCENTRATION GRADIENTS AND AEROBIC METABOLISM

The development of the geoelectric effect has been followedin Zea coleoptiles with a flowing-solution electrode system,and its dependence upon auxin concentration gradients and aerobicmetabolism assessed. A symmetrical source of IAA can effectively replace the coleoptiletip in allowing the geo-electric potential to occur. The diffusatefrom coleoptile tips, when applied asymmetrically to the apexof a vertical decapitated coleoptile, generates a potentialdifference across the coleoptile indistinguishable from thatinduced by the asymmetrical application of IAA. Asymmetricalapplication of IAA to vertical Avena and Zea coleoptiles andHelianthus hypocotyls induces closely similar responses. Neither the geoelectric effect nor a geotropic response developswhen intact Zea coleoptiles are placed horizontally after beingdeprived of oxygen, but they both occur when an aerobic atmosphereis restored. The lateral potential difference induced by theasymmetrical application of IAA to the apex of a vertical coleoptiledoes not occur under anoxic conditions. With a static-drop electrode system and a decapitated Zea coleoptile,a potential difference develops immediately after reorientationof the coleoptile into the horizontal position, and attainsa maximum value after about 10 min. This potential differencecan be further increased by the asymmetrical application ofIAA to the lower half of the apical cut surface of the coleoptile. Our data support the view that both the geoelectric potentialand the geotropic response are due to the IAA concentrationgradient which arises from the lateral transport of this substancefrom the upper to the lower half of the horizontal shoot. Theyalso bear out our previous conclusions that the ‘geoelectricpotential’ observed with static-drop electrodes and anintact shoot, is the resultant of two processes. The first isa physical phenomenon arising in the electrodes, or betweenthe electrodes and the plant tissue, and the second arises inthe living tissues of the shoot as the result of gravity-inducedchanges in auxin distribution.     

Studies in the Physiology of the Onion Plant: V. AN INVESTIGATION INTO THE GROWTH SUBSTANCE CONTENT OF BULBING ONIONS

Extracts have been prepared from induced and non-induced onionplants. These extracts have been fractionated and the ether-soluble,acidic components investigated using the wheat coleoptile straight-growthassay, colorimetric determination of 3-indolylacetic acid (IAA),and a new bioassay which has been introduced and modified togive reproducible results. This last is an assay of substanceswhich will cause ‘bulbing’ (swelling of the leafbases) of onion plants, and it has been used to demonstratea number of factors which influence the initiation of bulbingin the onion. It has been shown that substances are presentin extracts of onion tissue which will cause an increase inthe ‘bulbing ratio’ of test onion seedling sections. Results indicate that there is an increase in the IAA contentto a very high level during the first week following induction—beforethere is any visible sign of swelling of the leaf bases—butthat this falls off rapidly after 5–7 days and eventuallyfalls below the level found in non-induced plants.     

The Importance of Light Intensity for Pollen Tube Growth and Embryo Survival in Wheat x Maize Crosses

The success of Triticum aestivumxZea mays crosses, used to producewheat doubled haploids, is influenced by light intensity. Toexamine the basis for this response, pollen tube growth, embryosurvival and indicators of photosynthetic rate were measuredin two wheat cultivars (‘Karamu’ and ‘Kotuku’)crossed with maize at two irradiance levels (250 or 750 µmolm^-2s^-1, PAR). Pollen tube growth was significantly affectedby light intensity in ‘Karamu’ plants but not in‘Kotuku’ plants, despite both cultivars being pollinatedby the same maize source. The percentage of pollen tubes reachingthe cavity between the ovarian wall and integuments, or in themicropyle of ‘Karamu’ plants at high light intensity(65%) was nearly three-times greater than that at low lightintensity (22%). Thus, either low light intensity can affectthe maternal wheat plant in a way that inhibits pollen tubegrowth and/or high light intensity may promote pollen tube growthin ‘Karamu’ plants. Significant differences in ratesof electron transport in plants grown at the two light intensitiesindicated that the rate of photosynthesis may also have an effecton pollen tube growth. These results have importance for improvingthe efficiency of wheat x maize crosses and other wide cerealcrosses. Copyright 2001 Annals of Botany Company Intergeneric hybridization, light intensity, pollen tube growth, embryo survival, Triticum aestivum, wheat,Zea mays , maize     

Growth-regulator Interactions in the Growth of the Shoot System of Avena sativa Seedlings: I. THE GROWTH OF THE FIRST INTERNODE

1. Segments, 3.5 mm. long, cut from the first internode of Avenasativa seedlings grown in complete darkness respond to bothauxins and gibberellic acid by accelerated extension. 2. The optimum concentration of indole-3-acetic acid (IAA) is10 p.p.m. and of gibberellic acid (GA) is 0.1 p.p.m. 3. The degree of stimulation relative to the growth of controlsegments is affected by the inclusion in the segement of thenode between the internode and coleoptile. Thus the gibberellineffect is greatly increased while the IAA effect is decreased.The optimal concentrations are not affected by inclusion ofthe node. 4. These results can best be explained in terms of the supplyby the node tissue of an endogenous auxin which is necessaryfor the expression of GA action. 5. Numerous factorial experiments demonstrated that there isno detectable interaction between applied IAA and GA in thepromotion of first-internode extension. This implies that thepostulated endogenous auxin which synergized GAA action in (4)is either an active form of IAA produced only in the node tissueor is a completely different auxin. 6. No synergism of growth-promotive action can be detected betweenGA and the two synthetic auxins I-naphthylacetic acid and 2,4-dichlorophenoxyaceticacid. 7. p-chlorophenoxy-iso-butyric acid (PCIB) anc 2,4,6-trichlorophenoxyaceticacid (2,4,6-T) act as weak auxins and thus antagonize competitivelythe promotive action of GA. 8. The anti-auxin -(I-naphythyl-methyl-sulphide)propionic acid(NMSP) antagonizes competitively the promotive action of bothIAA and GA. 9. The facts under (5)–(8) suggest that auxins and GAare acting at the same growth-promotion centres and may competefor them. 10. Growth inhibitions are induced by high concentrations ofPCIB, 2,4,6-T and NMSP. The inhibitions produced by PCIB and2,4,6-T are both synergized by supra-optimal concentrationsof IAA while that of NMSP is synergized by supra-optimal concentrationsof both IAA and GA. This similarity of the effects of IAA andGA suggests that their inhibition actions also are of a closelysimilar nature.     

Growth Substances in the Roots of Vicia faba

An investigation has been made into the growth regulators presentin ethanol extracts of the seedling roots of Vicia faba afterseparation on paper partition chromatograms, using segmentsof Avena coleoptiles and mesocotyls and of Pisum sativum.rootsas biological assay material. Acetonitrile purification shows the presence of at least threeauxins running in isobutanol: methanol: water, at Rfs of 0–0·25,0·4–0·6, and 0·65–0·95;the latter may represent two different auxins. A similar, butclearer, picture is shown by the ether-soluble acid fraction.Here an auxin at Rf 0–0·25 also stimulates rootgrowth and could be ‘accelerator ’. A second atRf 0–0·25 is an indole compound which inhibitsroot growth and does not seem to be be IAA. A third at Rf 0·8–1·0is also a root-growth inhibitor and gives no indole reaction.The ‘inhibitor ß’ complex was demonstrated(Rf 0·65–0·85) together with a number ofother inhibitors at lower Rf value. The ether-soluble neutral component also contains auxins orauxin precursors. The water-soluble, ether-insoluble fraction contains four readilyinterconvertible substances with auxin properties. They allappear to inhibit root growth and give no indole reaction.     

Growth Substance Interactions during Uptake by Coleoptile Segments of Avena sativa and Hypocoty1 Segments of Phaseolus radiatus

Further studies have been made on the interactions of plant-growthregulators during uptake by Avena sativa coleoptile and Phaseolusradiatus hypocotyl segments. 2, 4-Dichlorophenoxyacetic acid(2, 4-D) had no effect on the uptake of either indol-3yl-aceticacid (IAA) or -naphthylacetic acid (NAA) by Avena. On the otherhand, a-(i-naphthylmethylthio)-propionic acid (NMSP) stronglyinhibited IAA uptake non-competitively but was much less effectiveon NAA uptake by Avena. The ‘metabolic’ uptake ofIAA by hypocotyl segments of Phaseolus radiatus was very stronglyinhibited by 2, 3, 5-tri-iodobenzoic acid (TIBA).     

Growth and Dormancy in Lunularia cruciata (L.) Dum: VII. THE ISOLATION AND BIOASSAY OF LUNULARIC ACID

The extraction, purification, and isolation of the growth inhibitorpreviously postulated are described. Methanol extraction andseparation into acid, neutral, and basic fractions was followedby paper chromatography of the acid and neutral fractions withdistilled water, re-extraction with methanol, and thin-layerchromatography, the peak of inhibition being located at Rf 0.7–0.8(isopropanol: ammonia: water, 100:5:5), or Rf 0.3–0.4(chloroform: ethyl acetate: acetic acid, 60:40:5) Lunularia gemmae, grown directly on the chromatographic stripwith added nutrient solution, served as the most appropriateand direct bioassay. Area measurements after 5–10 days'growth yielded significant differences. Other bioassays included:Marchantia polymorpha gemmae, lettuce hypocotyl growth, cress-seedgermination, oat coleoptile, and radish cotyledon disc tests.An active inhibitor, i.e. dihydrohydrangeic acid, now named‘lunularic acid’, was isolated in crystalline form.Lunularic acid was found to increase with long-day treatmentof Lunularia thalli, though present even in short-day. Its concentrationcould be altered rapidly when daylength conditions were changed.The growth inhibition was linearly related to concentrationover the range from 0.1 to 10 ppm, very high concentrationsbeing lethal. Abscisic acid, though inhibitory to Lunulariain low concentrations, was not detected in extracts, and couldeasily be separated from lunularic acid.     

MODE OF ACTION OF MALEIC HYDRAZIDE ON THE GROWTH OF ESCHERICHIA COLI AND AVENA COLEOPTILE SECTIONS

1. MH was found to suppress the growth and respiration of E. colias well as the IAA-induced growth of Avena coleoptile sections.
2. These suppressions could be reversed more or less strikinglyby the addition of a trace of heavy metals such as Co, Mn, Ni,Zn, Cu, or Mo.
3. The reversal could also be achieved by cysteine,thioglycollate,or fumarate, the latter two substances being,however, lesseffective.
4. The inhibition of the growth of E.coli by MH was completelyrelieved by the addition of IAA. Conversely,the inhibitionof the microbial growth by high concentrationsof IAA couldbe relieved by the addition of MH.
5. It was inferredthat MH may block certain heavy metal-catalyzedprocess, inwhich some thiol substance and IAA are participating,probablyby combining with the heavy metal.

(Received June 23, 1960; )     

Growth and Cellular Differentiation in the Wheat Coleoptile (Triticum vulgare L.): I. ESTIMATION OF CELL NUMBER, CELL VOLUME, AND CERTAIN NITROGENOUS CONSTITUENTS

The numbers and volumes of cells were determined for consecutivestages in the growth and development of the wheat coleoptile(var. ‘King II’) when grown at 25° C. in darknessfrom soon after germination to senescence. Cell expansion occurredthroughout growth and development up to 96 hours, and was accompaniedby cell division between 18–60 hours. Evidence is presentedthat suggests there are two phases of cell expansion concernedin coleoptile growth. Determinations were made at daily intervals from 24 to 120 hoursafter sowing of protein nitrogen, trichloroacetic acid solublenitrogen (TCA-sol. N), ribonucleic acid (RNA), and deoxyribonucleicacid (DNA) in the coleoptile, and the results expressed on aper-coleoptile and per-cell basis. The maximum rate of net proteinsynthesis took place during or after the cell-multiplicationphase of growth, depending on whether the results were expressedper coleoptile or per cell respectively. The ratio of proteinnitrogen to TCA-sol. N changed considerably during growth, from4·7 for young cells to 0·68 for mature cells. The fluctuations in the values for RNA and protein are consistentwith the template theory of protein synthesis and the DNA dataare discussed in relation to polyploidy in differentiated cells.No significant difference was found in the nucleotide compositionof RNA during the growth and development of the coleoptile.     

Studies on the Growth in Culture of Plant Cells: XI. THE INFLUENCE OF SHAKING RATE ON THE GROWTH OF SUSPENSION CULTUEES

The influence of shaking rates (expressed as revolutions permin) on orbital shaking platforms (1 in (2.54 cm) diam. rotarymotion) on the growth of cell suspension cultures of Acer pseudoplatanusL. and Atropa belladonna cultivar lutea Döll are described.By following cell growth and respiration and the levels of oxygenand carbon dioxide in the media during the progress of incubationit is concluded that the reduction of growth at sub-optimalshaking rates is not due to oxygen deficiency or toxic accumulationof carbon dioxide. The growth of the Atropa cell suspensionin ‘closed systems’ has been studied by the developmentof modified culture vessels and evidence obtained that the reducedgrowth in the systems is due to the formation by the culturesof an unidentified volatile growth inhibitor and not to eitheroxygen depletion or toxic accumulation of either carbon dioxideor ethylene. It is suggested that the reduced growth in ‘opensystems’ cultures at sub-optimal shaking speeds is eitherdue to retention of this volatile inhibitor or to restrictionof nutrient uptake by the existence of a stationary liquid-phaseboundary to the cells.     

The Galls on Forsythia intermedia Zab.

The galls on Forsythia intermedia Zab. consist of much-branched,root-like structures embedded in corky material which appearsto be sloughed-off from them. These gall ‘roots’have 7 to 14 vascular bundles compared with the 5 of normalroots and contain much more IAA. When galls were incubated in damp chambers the gall ‘roots’extended their growth. These extensions had a root-cap and root-hairsbut also had more vascular bundles than normal and they didnot respond geotropically like normal roots. Two fungi, Gibberella baccata (Wallr.) Sacc. (conid, stat. Fusarlumlateritium Nees.) and Phomopsis dominici Trav. were associatedwith die-back of gall ‘roots’ and two bacteria resemblingCorynebacterium fascians (Til.) Dowson and Agrobacterium tumefaciens(Sm. and Towns.) Conn respectively were isolated from galls.All produced IAA in culture media but their role in the etiologyof the gall, if any, remains in doubt.     

Studies in Stomatal Behaviour: VII. EFFECTS OF ANAEROBIC CONDITIONS UPON STOMATAL MOVEMENT--A TEST OF WILLIAMS'S HYPOTHESIS OF STOMATAL MECHANISM

An experiment was carried out to investigate stomatal responsesin wheat to four ‘closing treatments’, viz. highcarbon dioxide concentration, darkness, dry air and nil, eachgiven under both aerobic and anaerobic conditions. Thus theeffect of lack of oxygen on the closing (or opening) tendencywas estimated. Changes in calculated from resistance porometer readings were used as data and reasonsare given for thinking this is the best available measure forinvestigating stomatal dynamics in wheat. Williams's hypothesisdemands that lack of oxygen should cause stomatal opening orprevent closure; the present experiment shows that anaerobicconditions significantly increase the closing tendency when‘closing treatments’ are first applied. There isalso some suggestion that oxygen-lack itself tends to causeclosure in the absence of any other ‘closing treatment’.Williams's hypothesis in its original form is thus disproved(for wheat) but the present results would be consistent withan ‘active’ uptake of water by the guard cells contributingto stomatal opening. A nearly significant interaction betweencarbon dioxide and oxygen suggests that under anaerobic conditionsa ‘closing substance’ may perhaps be formed, forexample, by the union of some intermediate in glycolysis withcarbon dioxide.     

The Growth Substances separated from Plant Extracts by Chromatography: II. THE COLEOPTILE AND ROOT ELONGATION PROPERTIES OF THE GROWTH SUBSTANCES IN PLANT EXTRACTS

1. 1. A method for the running of ‘strip’ chromatogramsof plant extracts, as large-scale sources of the naturally occurringgrowth substances accelerator () and inhibitor ß(ß), and the elution of these substances togetherwith indole-3-acetic acid (IAA), is described. A method is givenfor the testing of the pea root section extension propertiesof these growth substances.
2. 2. Coleoptile and root sectionextension tests over a completeconcentration range are donefor , ß, and eluted IAA,and mixtures of and ßwith IAA or indole-3-acetonitrile(IAN) are tested for coleoptilesection extension.
3. 3. promotes at low concentrations andinhibits at high concentrationsboth coleoptile and root sectionextension and the coleoptilesection extension induced by IAAor IAN. ß inhibitscoleoptile and root section extensionover the whole concentrationrange; it also inhibits IAA andIAN induced coleoptile sectionextension.
4. 4. The extensionof coleoptile sections in mixtures of or ßwith IAAis measured at a number of time intervals. , aloneand withIAA, has its greatest promoting effect in the earlystages andits greatest inhibiting effect in the later stagesof sectiongrowth. ß, alone, promotes the early stagesand inhibitsthe later stages of section growth and, with IAA,has its greatestinhibitory effects in the later stages.