An Investigation of the Relay Hypothesis of Phloem Transport in Ricinus communis L.

To test for the existence of an apoplastic unloading/reloadingstep in phloem translocation, as envisaged in the relay hypothesisof phloem transport, isotopic trapping experiments were performedon Ricinus comunis L. var. Gibsonii. A CO₂ buffer system wasused to supply ¹⁴CO₂ at constant partial pressure and constantspecific activity to a photosynthesizing leaf. The subtendinginternode was perfused with solutions of sugars or of mannitoland transolation of ¹⁴C past the perfused zone was monitoredby the collection of phloem exudate. Trapping of activity inthe perfusate was enhanced by the presence of sugars as wouldbe expected with an unloading/reloading process. However, therewas no evidence that introduction of the unlabelled sugars tothe apoplast also reduced activity in the phloem exudate. Moreover,the rate of loss of activity to the perfusing solutions representedonly 1% of the rate of appearance in the exudation. It is suggestedthat the trapping results may reflect an unloading of tracerfrom the phloem associated with a subsequent reloading by adjacenttissues rather than by the sieve tubes. To investigate the length of sieve tube continuity in Ricinus,a horizontal incision was made to the bark and the rate of exudationof phloem sap was monitored. Successive circumferential cutswere made above the exuding incision and progressively closerto it. In general, a girdling incision produced a transientdecrease in the rate of change of exudation rate (i.e. the firstderivative became more negative/less positive). The magnitudeof this response rose with exudation rate and fell with thedistance at which a girdling cut was made. Fitting an appropriatemodel yielded an estimate for contributory length of 69 ±6 cm. This was comparable with the distance of the initial tangentialincision from the stem apex, suggesting a continuous sieve tubesystem in Ricinus. A similar investigation on the petiole yieldedan estimate of around 7.0 cm. This lower estimate for contributorylength is believed to reflect a rapid sealing process that limitsthe distance of propagation of turgor-release rather than alimited length of sieve tube continuity. The results of this investigation do not support the relay hypothesisof phloem transport. Rather they suggest a continuous sievetube system which has a distributed capacity to load and unloadsolutes, and which may exhibit a sealing response when injured. Key words: Ricinus communis L, phloem transport, phleom unloading     

Studies on the Movement of Solutes between the Sieve Tubes and Surrounding Tissues4: II. PATHWAYS OF ION TRANSPORT FROM THE XYLEM TO THE PHLOEM

It has been shown that the time lapse of 2–3 hours foundby Peel (1963), between the introduction of an inactive ioninto the xylem of a stem segment and its appearance in honeydewcollected from that segment, is not an artefact caused by detectionmethods. A delay of similar magnitude was found in the presentexperiments when radioactive rubidium, ³²P-phosphate, or ³⁵S-sulphatewas used. Pretreatment of stem segments with the respectiveinactive ion caused a very considerable increase (20 hours)in the time lapse when the radioactive ions were subsequentlysupplied to the segment. Experiments with bark strips demonstratedthat the time lapse was due to the presence of a barrier tothe radial movement of ions from the xylem of segments, sinceusing bark strips the time lapse was reduced to less than onehour. This barrier could either be the xylem itself, the cambium,or both tissues. Pretreatment of bark stnps with inactive ionsincreased the time lapse in the case of ³²P-phosphate and ³⁵Ssulphate, but had no detectable effect with radioactive rubidium.It is considered that these results indicate that rubidium maybe able to pass directly into the sieve tube from the bathingsolution in bark strips. Phosphate and sulphate, however, mayhave to move into the sieve tube via the storage cells of thebark. It has been shown that phosphorus-32 activity in the sievetube is not confined to inorganic phosphate, but is certainlypresent in other compounds of which three sugar phosphates havebeen identified.     

A Reanalysis of Particle Motion in Sieve Tubes of Heracleum

From an analysis of the Brownian motion of particles in sieveelements of Heracleum mantegazzianum and Heracleum sphondylium,Barclay and Johnson have suggested that the in situ viscosityof sieve tube sap is four to six times higher than has previouslybeen assumed. In particular, they obtained a value for the sapviscosity of about 10^–2 Pa s, which compares with a valueof 2 x 10^–3 Pa s for a 20 per cent (w/v) sucrose solution.The present paper describes a reanalysis of their data. It isargued that Barclay & Johnson underestimated the Brownianmotion of sieve element particles and so overestimated the sapviscosity. An exact correction was not possible, but it is concludedthat the in situ viscosity of Heracleum sieve tube sap mustbe less than 3 x 10^–3 Pa s, which corresponds to a sucroseconcentration of less than 29 per cent. Hence it may not beunreasonable to suppose that the viscosity of sieve tube sapis determined primarily by the concentration of sucrose, ashas been assumed in theoretical analyses of the Munch hypothesis.It is also concluded that the sieve tubes studied by Barclayand Johnson were not functional, in the sense that they didnot exhibit an axial bulk flow of sap. Heracleum, sieve tubes, Brownian motion, viscosity, Munch hypothesis     

The Physiology of the Sieve Tube: PRELIMINARY EXPERIMENTS USING APHID MOUTH PARTS

The willow aphid Tuberolachnus salignus feeds by inserting itsstylets into a single sieve element. If the insect is severedfrom its proboscis whilst feeding, sap from the sieve tube exudesfrom the cut end of the stylet canal and may continue for manyhours. By attaching a capillary tube the rate of exudation canbe measured and the sap analysed. Normally the rate of exudationwas about 1 mm.³ per h. This means a movement of water and sucroseof 100 cm./h. or 100 sieve elements per min. The sap was foundto contain between 5 and 15 per cent, sucrose, up to 0.4 percent, raffinose, no reducing sugars, and about 0.5 per cent,amino-acids. Girdles some distance above and below exuding stylets failedto stop exudation (Tables I and II). This indicates a rapidsealing of the cut ends of the sieve tube and a switch-overin source of supply from the leaves to storage cells in thestem. The use of isolated stem segments and even irrigated stripsof bark followed. Using these, the following points have beenestablished, (a) There is no polarity of movement (Table IV);(b) a minimal length of about 16 cm. of stem or 800–1,000sieve elements are needed to give the full exudation rate (Fig.5); (c) when the D.P.D. on the inside of the bark is raised(Figs. 6, 7, and 8), the rate of exudation declines, but theconcentration of the exudate rises and exudation continues,indicating a maintenance of turgor, even in face of a D.P.D.of 20 atm. or more. Secretion of sugar into the sieve tube continuesalbeit slowly even against a sieve tube concentration of 50per cent. These findings are discussed in relation to current theoriesof translocation and are considered to favour the mass flowhypothesis.     

Investigation of Bidirectional Movement of Tracers in Sieve Tubes of Salix Viminalis L.

Sieve-tube sap was collected as honeydew from the aphid Tuberolachnussalignus (Gmelin) feeding on a young shoot or a mature stemof willow (Salix viminalis L.). Two radioactive tracers (either³H-glucose and ¹⁴C-sucrose, or ¹⁴CO₂, Na₂ ¹⁴CO₃, and ³²P-orthophosphate)were applied to leaves or to bark abrasions at each end of thestem. In some experiments the tracers were applied after thestart of honeydew production, whilst in others the tracers wereapplied before the aphids were sited. In most cases aphids feeding between the tracer applicationpoints produced doubly-labelled honeydew during a 24–48-hperiod from tracer application. In some instances the firstdrops of honeydew contained one tracer, followed several hourslater by the other tracer, whilst some aphids produced doubly-labelledhoneydew from the first drop. Singly-labelled or inactive honeydewwas found in only a few cases. It was demonstrated that when ³H-glucose and ¹⁴C-sucrose wereapplied at opposite ends of a stem that the tracers moved inopposite directions in the form of sucrose. The data suggest that a simultaneous bidirectional movementof two tracers may occur in the same sieve tube. However, thepossibility that the doubly-labelled honeydew could be producedby lateral movement from one sieve tube carrying one tracerto an adjacent punctured sieve tube carrying the other tracercannot be discounted.     

The Effect of Calcium Starvation on Assimilate Partitioning and Mineral Distribution of the Phloem

Populus plants were grown in a medium lacking calcium and exposedto ¹⁴CO₂. In contrast to plants in the complete nutrient medium,the percentage amount of ¹⁴C-assimilates increased in the leavesof calcium-deficient plants and decreased in the stem and theroots. When plants were grown without potassium or magnesiumno differences in the amount of ¹⁴C-label occurred in comparisonwith plants in the complete nutrient medium. Translocation wasrecorded by microautoradiography. It was observed that considerableamounts of labelled photoassimilates were unloaded from thephloem in the middle part of the stem in plants of the completenutrient medium. In contrast, during calcium starvation ¹⁴C-labelwas restricted to the phloem of the stem. In addition, the concentrationsof magnesium and phosphorus showed a remarkable increase instem sieve tubes of calcium-deficient plants. When sieve tubesof source leaves from Populus, barley and maize were comparedwith those of sink leaves, the latter showed higher calciumconcentrations. The results suggest that calcium is a necessaryfactor in the regulation of phloem translocation. Key words: Calcium deficiency, phloem translocation, sieve element loading and unloading, X-ray microanalysis     

Studies on the Movement of Solutes between the Sieve Tubes and Surrounding Tissues3: I. INTERFERENCE BETWEEN SOLUTES AND RATE OF TRANSLOCATION MEASUREMENTS

Certain aspects of the secretion of solutes into, and removalfrom, the sieve tubes of isolated stem segments and rooted cuttingsof Salix viminalis have been studied. Sieve-tube sap was obtainedeither as honeydew from whole individuals or via the severedstylets of the aphid Tuberolachnus salignus (Gmelin). It was shown that interference occurred between the chemicallyunrelated solutes, sucrose and the cations potassium and rubidium.On raising the potassium concentration in the sieve-tube sapby passing a solution of this ion through the xylem, the sucroseconcentration declined. When the sucrose concentration fellover a period of days due to respiratory loss of carbohydratesfrom an isolated stem segment, a concomitant rise in eitherthe potassium or rubidium level in the sap occurred. When a solution of sodium was passed through the xylem, theconcentration of this ion in the sieve-tube sap rose, whilstthat of potassium fell at first, but later rose higher thanits initial value, indicating that both antagonism and synergycan occur between these ions. On introducing both these cationsinto the xylem simultaneously, more sodium than potassium wastaken up by the segment, though the increase in the sodium concentrationin the sieve-tube sap was less than that of the potassium. Perfusingthe xylem with a calcium solution had no effect upon the concentrationof potassium in the sieve tube. It has been shown that the rate of translocation of a solutealong the sieve tube, as measured by the two colony technique,depends upon the rate of removal of this solute from the sievetube. The amount of such lateral loss from the sieve tube isrelated to the potential gradient for a solute between the sievetube and surrounding cells.     

The Effect of Changes in the Diffusion Potential of the Xylem Water on Sieve-tube Exudation from Isolated Stem Segments

¹⁴CO₂, was supplied to rooted leafy cuttings of Salix viminalis.Several hours after activity had reached a point on the stem50 cm below the leafy shoots (indicated by activity being detectablein honeydew collected from a colony of the aphid Tuberolachnussalignus) the stem was severed at this point, and also some10 cm below the shoots. This gave a segment approximately 40cm in length, in which there was a gradient of ¹⁴C activity,this being highest at the apical end, and lowest at the basalend. An aphid colony was then sited at the low-activity endof the segment and the effect of changes in the diffusion potentialof the xylem water on the specific activity of honeydew fromthis colony was investigated. It was shown that gradients ofhydrostatic pressure affected the specific activity. With thecolony at the low-pressure end of the segment the specific activityincreased, the converse being found with the colony at the high-pressureend. Uniform changes in diffusion potential also produced changesin specific activity: an increase in hydrostatic pressure causinga rise, and a lowering of the diffusion potential, using anosmoticum, a fall. These changes in specific activity are interpretedas being due to changes in the length of segment supplying solutesto the aphid colony.     

The Cellular Pathway of Short-distance Transfer of Photosynthates and Potassium in the Elongating Stem ofPhaseolus vulgarisL. A Physiological Assessment

Plasmolytic disruption of plasmodesmata interconnecting metaphloemsieve element-companion cell complexes with small and largephloem parenchyma cells in the elongating region of internode2 ofPhaseolus vulgarisL. seedlings did not affect accumulationof phloem-imported¹⁴C-photosynthates and⁸⁶rubidium. The membrane-impermeantdye, 5(6) carboxyfluorescein, loaded into leaf phloem as themembrane-permeant diacetate ester, was found not to move radiallyout of the importing sieve elements in the internode elongationregion. In contrast, the apoplasmic tracer, Calcuofluor White,rapidly moved laterally throughout all tissues of the elongationzone. Hexoses, sucrose and potassium were identified as themain osmotica in internode apoplasmic sap. Label asymmetry in[¹⁴C](fructosyl)sucrose was retained on accumulation by excisedstem segments. Uptake of [¹⁴C]sucrose and⁸⁶rubidium by stemsegments exhibited saturation kinetics. Sucrose uptake was inhibitedby the slowly penetrating sulphydryl reagent, para-chloromercuribenzenesulphonicacid.In vitrorates of sucrose uptake, at apoplasmic concentrations,corresponded to its predictedin vivorate of delivery to thestem ground tissues from mature sieve elements when respiratorylosses were assumed to be confined to the stem phloem. For potassium,the total delivery rate could be accounted for by itsin vitrorateof uptake. Overall, it was concluded that radial transport,in the elongation zone of internode 2 ofPhaseolus vulgarisL.seedlings, follows an apoplasmic route from mature sieve elementsto stem ground tissues.Copyright 1998 Annals of Botany Company PhaseoluLes vulgaris, apoplasm, elongating stem, French bean, photosynthates, potassium, radial transfer, symplasm.     

Auxin-promoted Transport of Metabolites in Stems of Phaseolus vulgaris L: EFFECTS AT THE SITE OF HORMONE APPLICATION

The hypothesis that indol-3yl-acetic acid (IAA) stimulates solutemobilization in stems by increasing metabolite utilization (i.e.sink strength) and/or sieve tube unloading mechanisms was investigatedusing seedlings of Phaseolus vulgaris L. Sink strength was assessedby obtaining estimates of rates of growth, ¹⁴C incorporationinto protein, sucrose metabolism, and sucrose uptake. The dataobtained suggest that, whilst enhancing assimilate mobilization,IAA had no short term effect on the sink strength of the treatedstumps. In several experiments involving long distance transportof ¹⁴C-assimilates, the magnitude of auxin-promoted transportwas found to be insensitive to potential changes in sink strength.Furthermore, ¹⁴C transport profiles demonstrated that the siteof hormone action was not confined to its point of application.     

A Reanalysis of the Kinetics of 14C Photosynthate Translocation in Morning Glory Vines

The spatial and temporal profiles of ¹⁴C photosynthate observedafter a 5 min pulse-labelling period in morning glory vinesare examined. In these experiments a peak of radioactivity appearsto travel down the stem, growing shorter and broader with timeand distance. It is possible to fit the known data reasonably well with aHorwitz reversible-loss model in which the observed changesin peak height and width are attributed to the reversible exchangeof tracer molecules between the translocating sieve tube anda stationary reservoir. The parameters that determine the spatialand temporal profiles are the input kinetics, the sap velocity,v, the ratio of the reservoir volume to sieve tube volume, ,and the permeability to radius ratio, P/r. Even when the inputkinetics are known it is possible to obtain several equallygood fits to the observed tracer profiles with different combinationsof v, , and P/r. With our present state of knowledge it is notpossible to characterize these parameters uniquely, and it isunlikely that tracer techniques alone will yield definite valuesfor v, , and P/r. Ipomoea oil Roth, morning glory, translocation profiles, Horwitz reversible-loss model, mathematical models     

Respiration and Phloem Translocation in the Roots of Chickpea (Cicer arietinum)

Root growth in chickpea (Cicer arietinum) has been studied fromthe early vegetative phase to the reproductive stage in orderto elucidate its growth and maintenance respiration and to quantifythe translocation of assimilates from shoot to root. A carbonbalance has been drawn for this purpose using the growth andrespiration data. The increase in the sieve tube cross-sectionalarea was also followed simultaneously. Plants growing in a nutrient culture medium were studied todetermine the relative growth rate (RGR) 5–60 d aftergermination. RGR declined from 113 to 41 mg d^–1 g^–1during the measurement period. Simultaneous with the RGR analysis,respiration rate was also measured using an oxygen electrode.The respiration rate declined as the plants aged and a drasticreduction was recorded following anthesis. The relationshipbetween RGR and respiration rate was used to extrapolate themaintenance respiration (m) and growth respiration (1/Y_EG).The respiration quotient (r.q.) of the roots was 1.2 and theQ₁₀ in the range 20–25 °C was 2·2. A carbon balance for the roots was constructed by subtractingthe carbon lost during respiration from that gained during growth.The roots were found to respire no less than 80% of the carbontranslocated. The increase in the cross-sectional area composed of sieve tubeswas measured near the root-shoot junction as the plants grew.Chickpea has storied sieve plates which simplifies these measurements.Their cross-sectional area increased during growth mainly becauseof an increase in sieve tube number. The diameter of individualsieve tubes remained constant. Specific mass transfer (SMT) values for seive tubes into theroots have been computed during various stages of growth. SMTvalues were relatively constant before anthesis (approx. 6·5g h^–1 cm^–2), but decreased following anthesis. Wedid not evaluate possible retranslocation from roots: any suchretranslocation would have the effect of increasing our SMTvalues. Chickpea, Cicer arietinum, legume, root, respiration, phloem, translocation, carbon balance, specific mass transfer, sieve-tube dimensions     

Transport of 14C-IAA in Cucurbita maxima: A Comparative Study of Rapid, Non-Polar Movement and Slow, Polar Auxin Transport

The transport of ¹⁴C-IAA has been studied in Cucurbita maxima.IAA fed to the leaf of an intact plant moves rapidly in a non-polarfashion in the phloem. Collection and analysis of exudate fromsevered sieve tubes showed that there was no metabolic conversionor complexing of IAA for several hours. Polar movement of ¹⁴C-IAA in isolated internode segments occursat rates an order of magnitude slower than movement in the phloem.The importance of discrete and isolated channels of hormonetransport, that vary in direction and rates, is briefly discussed.     

A Working Model of a Sieve Tube

A working model of a sieve tube is described, based upon thePressure-Flow Hypothesis. The flow of solution in the sievetube is envisaged as being due to the joint influence of anaxial turgor pressure gradient and a lateral water potentialdifference which causes an intake of water along the tube. Inthis second respect the model differs from that originally suggestedby Münch. The model comprises a length of dialysis tubing rendered semi-permeable,incorporating capillary resistances at regular intervals. Asucrose solution is pumped into one end and collected at theother, the whole being submerged in water. Turgor pressures and concentrations along the model were recordedat intervals; thus the approach to the steady state was followed.In the steady state ¹⁴C-sucrose was introduced to the solutionbeing pumped and its approach to a steady distribution in themodel was followed. Important conclusions reached are that the Munch Pressure-FlowHypothesis implies a pressure profile convex upwards, a velocityincrease down the phloem, and an exponential fall in tracerconcentration without its lateral leakage.     

The Cellular Pathway of Radial Transfer of Photosynthates in Stems of Phaseolus vulgaris L.: Effects of Cellular Plasmolysis and p-Chloromercuribenzene Sulphonic Acid

Cellular plasmolysis with l M solutions of mannitol appearedto sever plasmodesmatal interconnections between all cells ofthe stems of Phaseolus vulgaris plants except the sieve element-companioncell (se—cc) complexes. Phloem loading and uptake of [¹⁴C]sucroseby the storage cells of the stems was unimpaired by cellularplasmolysis followed by rehydration of the stem tissues. Accumulationof phloem-transported ¹⁴C-photosynthates of the treated stemswas inhibited in summer-grown plants and unaffected in winter-grownplants indicating that phloem unloading follows a symplasticand a free-space route respectively depending on growth season.At a concentration that did not interfere with cellular metabolism,p-chloromercuribenzene sulphonic acid (PCMBS) applied to thestems blocked [¹⁴C]sucrose loading into the phloem and storagecells of the stem, but had no effect on the pool size of free-spacesugars. This latter response is consistent with a facilitatedmechanism of sugar unloading to the stem free-space. Accumulationof phloem-transported ¹⁴C-photosynthates was stimulated by PCMBSand this effect was most pronounced in winter-grown plants.Cellular plasmolysis followed by rehydration abolished the PCMBSaction on ¹⁴C-photosynthate accumulation. This effect is consistentwith a PCMBS induction of phloem unloading through the stemsymplast. It is proposed that phloem unloading in bean stemsmay follow either a free-space or symplastic route and thatthe latter route is entrained under sink-limited conditions. Phaseolus vulgaris, french bean, stem, phioem unloading, free-space, symplast     

Further Evidence of Apoplastic Unloading into the Stem of Bean: Identification of the Phloem Buffering Pool

Further evidence that sucrose is passively leaked from the sievetubes directly into the apoplast of bean stems and activelyreloaded is reported. The stem apoplast is identified as thepool of sucrose outside the sieve tubes which buffers againstsudden changes in sieve tube sucrose concentration, caused bysudden changes in sink demand or source supply. Key words: Sucrose transport, Phaseolus vulgaris, Apoplast, Phloem unloading     

Cell Length Variation in the Secondary Phloem of Dalbergia spp. with Increasing Age of the Vascular Cambium

Cell length variation of sieve tube elements and phloem fibreswas studied in relation to their relative position within thebark in five species of Dalbergia. It was found that in fourout of the five species studied, the fibre elements occurringin the vicinity of cambium are longer than those occurring atthe periphery of the bark. D. latifolia is found to be an exception,since the shorter elements occur near the cambium while thelonger elements are found in the outer regions of the bark inthis species. In all the species fibre length declines markedlyin the middle region of the bark, and this is followed by asudden but temporary rise, except in D. latifolia in which theshortest elements occur both in the middle as well as near thecambium. The cell length variation of sieve tube cells does not followany recognizable trend in any of the species studied.     

The Pathway of Radial Transfer of Photosynthate in Decapitated Stems of Phaseolus vulgaris L.

[¹⁴C]Sucrose was found to be the predominant component of the¹⁴C-photosynthates that accumulated in the free space of decapitatedstems of P. vulgaris plants. The ¹⁴C-photosynthates appearedto occupy the entire free-space volume of the stems at totalsugar concentrations in the range of 3–12 mM. The free-spacesugar levels were found to rapidly decline once photosynthatetransfer to the stems was halted. Moreover, it was found thatestimates of the rate of in vitro sucrose uptake by the stemscould account fully for the decline in free-space sugar levels.Overall, the evidence indicated that at least part of the radialpathway of photosynthate transfer in bean stems involved thestem apoplast. It is tentatively proposed that, based on celland tissue distribution of ¹⁴C-photosynthates, the apoplasticpathway extends from the membrane boundary of the sieve element/companion-cellcomplex to all other cells of the stem. Apoplast, Phaseolus vulgaris L., bean, phloem unloading, photosynthates, symplast     

Auxin-Promoted Transport of Metabolites in Stems of Phaseolus vulgaris L.: Auxin Dose-Response Curves and Effects of Inhibitors of Polar Auxin Transport

Transport of ¹⁴C-photosynthate in decapitated stems of Phaseolusvulgaris explants was dependent on the concentration of indole-3-aceticacid (IAA) applied to the cut surfaces of the stem stumps. Thephysiological age of the stem influenced the nature of the transportresponse to IAA with stems that had ceased elongation exhibitinga more pronounced response with a distinct optimum. Increasednutrient status of the explants had little influence on theshape of the IAA dose-response curve but increased, by two ordersof magnitude, the IAA concentration that elicited the optimalresponse. Applications of the inhibitor of polar auxin transport,1-(2-carboxyphenyl)-3-phenylpropane-1, 3-dione (CPD), affectedIAA-promoted transport of ¹⁴C-photosynthates. At sub-optimalIAA concentrations, CPD inhibited transport, whereas at supra-optimalIAA concentrations, ¹⁴C-photosynthate transport was marginallystimulated by CPD. Treatment with CPD resulted in a significantreduction in stem levels of [¹⁴C]IAA below the site of inhibitorapplication, while above this point, levels of [¹⁴C]1AA remainedunaltered. The divergent responses of auxin-promoted transportto CPD treatment are most consistent with a remote action ofIAA on photosynthate transport in the decapitated stems. Key words: Auxin, photosynthate, transport     

The Uptake of Growth Substances: XIII. DIFFERENTIAL UPTAKE OF INDOL-3YL-ACETIC ACID THROUGH THE EPIDERMAL AND CUT SURFACES OF ETIOLATED STEM SEGMENTS

The patterns of uptake of indol-3yl-acetic acid (IAA-2-¹⁴C)by etiolated stem segments of varying lengths have been examined,employing tissues excised from (a) the first and third internodesof Pisum sativum, (b) the top and base of the hypocotyl of Gossypiumhirsutum, and (c) the mesocotyl of Avena sativa. For all species,concentrations (10^–5–10^–3 M) and times upto 24 h, there is a steady accumulation of radioactivity inthe segments. For equal volumes of tissue uptake is inverselycorrelated with segment length but for extending tissues theinitial enhanced extension growth is independent of length;that is there is no direct linkage between the rate of extensionand auxin content. Comparisons between segments with free andsealed ends established that over 24 h some 57–73 percent of the IAA enters via the cut surfaces. Initially, thepercentage is greater; expressed as a rate per unit of surfacethe differences between cut and epidermal surfaces can reach28-fold. The rate of entry through the epidermal surface islinearly proportional to the external concentration but thisdoes not hold for cut surfaces. The addition of streptomycin,synthalin, cetyltrimethylammoniumbromide (CTAB), and chitosanto the external medium does not promote uptake of IAA by Pisumsegments; indeed synthalin is markedly inhibitory. With Gossypiumsynthalin causes little inhibition. Larger depressive effectswere induced for entry via the cut surfaces. On entry the IAAis rapidly metabolized and the rate of conversion is higherfor segments with sealed ends. These findings are discussedin relation to (a) differences in the mechanisms determiningthe uptake of IAA and other auxins, (b) cell extension and thedistribution of auxin in the tissues.