Control of root growth: effects of carbohydrates on the extension, branching and rate of respiration of different fractions of wheat roots

Rates of extension, numbers of laterals and rates of respiration were measured in different fractions of wheat ( Triticum aestivum L. cv. Alexandria) roots following changes in carbohydrate supply. The supply of carbohydrate was varied by selective pruning and exogenously fed sugars. Pruning shoots to a single leaf (leaf-pruning) reduced the rate of O₂ uptake by intact roots. Rates were not stimulated by shortterm feeding of sucrose (25 m M ), but were stimulated by the uncoupler p -trifluoro-methoxy(carbonylcyanide)phenylhydrazone (FCCP). Feeding glucose to roots of leaf-pruned and non-pruned plants for 16–24 h increased the rate of O₂ uptake. It is concluded that respiration is under fine control by adenylates and coarse control by carbohydrate supply, with carbohydrates regulating directly the rate of some energy consuming process(es). These energy consuming processes are located in growing tissue fractions. Feeding glucose to leaf-pruned and non-pruned plants increased rates of O₂ uptake in seminal root tips, the zone of developing lateral primordia and mature root sections with elongating laterals, but had no effect on mature sections from which the laterals had been excised. Leaf-pruning reduced the extension rate of seminal axes and first-order laterals when measured over 24 h. Feeding glucose to roots from the time of pruning increased the rate, but did not fully restore it to control values. Pruning roots to a single seminal axis (root-pruning) and feeding glucose to non-pruned plants had no effect on the extension rate of the seminal axis or its laterals over this time period, although rates were increased by root-pruning when measured over 3 days. The number of lateral root primordia was reduced by leaf-pruning and increased by root-pruning and feeding glucose. The results are discussed in terms of the role of carbohydrates in the control of root growth and branching.     

The occurrence of free and bound cytokinins in clubroots and Plasmodiophora brassicae infected turnip tissue cultures

This paper deals with the quantitative determination of free and bound cytokinins in clubroot tissue and in Plasmodiophora brassicae Woron, infected Brassica campestris L. callus tissue. The fractions were separated in a butanol soluble fraction containing the free cytokinins such as zeatin and zeatin riboside and a water soluble fraction containing the bound cytokinins. The butanol fraction was extensively purified and analysed by high pressure liquid chromatography (HPLC). The butanol fraction contained cytokinins which cochromatographed with zeatin and zeatin riboside and not with dihydrozeatin. Zeatin and zeatin riboside were quantitatively determined by HPLC. Recovery of the cytokinins varied between 30–50%. Clubs contained 50–160 ng zeatin and 210–300 ng zeatin riboside per g dry weight. Callus tissue contained 133 ng zeatin and 169 ng zeatin riboside per g dry weight. Clubs, callus as well as healthy tissue contain large amounts of bound cytokinins. Upon treatment of the water soluble fraction first with alkaline phosphatase and then with β-glucosidase biologically active fractions were found which coeluted with zeatin and zeatin riboside on Sephadex LH₂₀ in 20% ethanol. Evidence is presented for a novel cytokinin in the water soluble fraction which yields free zeatin and glucose-6-phosphate after treatment with β-glucosidase.     

Extension rate and respiratory activity in the growth zone of wheat roots: Time-course for adjustments after defoliation

The time-course for adjustments in the rate of extension of wheat (Triticum aestivum L. cv. Alexandria) roots, and the activity and capacity of respiratory pathways in the root apex, were determined after pruning the shoot to the ligule of the first leaf. Leaf pruning reduced the extension rate of both seminal and lateral roots. The onset of the response occurred within 1 h of pruning for laterals and between 2 and 3 h for seminals. The reduction in rate appears to be the result of a decrease in carbohydrate availability because (1) in seminal roots it was preceded by a decrease in soluble sugar content of the apical part of the growth zone (0–5 mm behind the root apex) and (2) supplying glucose (50 mM) to the roots of plants defoliated 24 h earlier led to a steady increase in extension rate of both seminal and lateral roots compared to non-fed controls. Supplying 3-O-methyl glucose had no effect. The reduction in extension rate of seminal roots was accompanied (or slightly preceded) by a reduction in respiratory O₂ uptake in the apical part of the growth zone (0–5 mm). Changes in respiratory activity in the basal part of the growth zone (5–10 mm) only occurred several hours later. At the time root extension rate was reduced, the rate of O₂ uptake could be stimulated with FCCP, which indicates that respiration was under the fine control of adenylates. From these results we suggest the following sequence of events occurs after defoliation. Firstly, defoliation reduces the supply of sugars to the root apex, this leads to a reduction in rate of extension through some form of coarse control by carbohydrates on cell division and expansion, which in turn reduces the rate of respiratory O₂ uptake because of a smaller demand for ATP. The results also indicate that there is a rapid (<1.5 h) reduction in respiratory capacity in the root apex after defoliation which occurs before any change in the overall rate of respiration.     

Pathogenesis of barley leaves by Helminthosporium teres I: Green island formation and the possible involvement of cytokinins

Infection sites/green islands were formed in host leaf tissue infected with drops of H. teres. They exhibited higher cytokinin-like activity, sugar and starch than their surrounding tissue and tissue under water drops. The cytokinin-like activity at the infection sites increased from 24 to 72 h of incubation. However, the cytokinin-like activity of the tissue surrounding the infection drops and the tissue under water drops fell from 24 to 72 h incubation. The culture filtrate extracts of the fungus also produced cytokinin-like activity which increased from 1 to 10 days incubation. Application of this culture filtrate extract evoked green island formation. Application of kinetin to host leaves duplicated the green island effect. Thin-layer chromatographic fractions of the tissue extracts and the culture filtrate extracts revealed that a major portion of cytokinin-like activity corresponded to zeatin and zeatin riboside. The presence of zeatin and zeatin riboside (both in tissue and culture filtrate extracts) was confirmed by high performance liquid chromatography. Increases in the amounts of cytokinin-like substances, particularly zeatin and zeatin riboside, attributed to pathogen influence are suggested to be involved in infection and pathogenicity of H. teres.     

The effect of zeatin and iso-pentenyladenine on IAA transport from the shoot to the root of Pinus pinea seedlings

The tips of the tap roots of Pinus pinea seedlings were dipped in zeatin or iso-pentenyladenine solutions. Immediately after cytokinin application to the root tip or after a 24 h lag phase, [2-¹⁴C]IAA was applied to the shoot apex. Treating with zeatin resulted in an increase in [2-¹⁴C]IAA transport from the shoot to the root. Iso-pentenyladenine also caused a slight increase in transport of radioactivity to the root but this was less pronounced compared to the results obtained with zeatin. With zeatin treatment increasing amounts of radioactivity accumulated in the lateral root emerging zone of the tap root (Section III). This was in sharp contrast to the treatment with iso-pentenyladenine where little radioactivity accumulated in this section of the root. Recovery of radioactivity 48 h after applying [2-¹⁴C]IAA showed that 33% of the recovered radioactivity co-chromatographed with authentic IAA. The implications of the effect of different cytokinins on the distribution of radioactivity along the tap root of Pinus pinea following [2-¹⁴C]IAA application to the shoot are discussed.Abbreviations Z zeatin - iP iso-pentenyladenine - TCL thin-layer chromatography     

Cytokinin Activity in Lupinus albus

The cytokinin content of the root exudate and leaves of fruiting white lupin plants (Lupinus albus L.) was investigated at 2 weekly intervals after anthesis of the lowest flower on the primary inflorescence. Up to 8 weeks after anthesis the level of cytokinins in the root exudate increased. However, at 10 weeks after anthesis insufficient sap was produced for analysis. Cytokinins co-eluting with zeatin and zeatin riboside were detected in the root exudate after fractionation on Sephadex LH-20. The cytokinin levels in the mature leaves steadily increased up to 8 weeks after anthesis and thereafter remained relatively constant. Three peaks of activity, co-eluting with zeatin, zeatin riboside and the glucoside cytokinins were recorded in the leaf extracts. The level of glucoside cytokinins in the leaves was high at 8 and 10 weeks after anthesis. Paper chromatography of extracts of fruits collected at 2 weeks after anthesis indicated that as fruit development proceeded there was a build up of cytokinin in this region of the plant. It is suggested that, in the white lupin, the cytokinins translocated to the shoot are accumulated in the leaves and in the fruits and that it is only later when there is a considerable decrease in sap (10 weeks after anthesis) production that a decreasing supply of cytokinins leads to shoot senescence.     

The Effect of Ringing on Cytokinin Distribution in Salix baylonica

Although quantitative differences were observed in the cytokinin content of mature leaves and bark of Salix babylonica it would appear as if these tissues contained the same cytokinin complement. Ringing resulted in a decrease in the level of cytokinins in the leaves and an increase in the bark, both above and below the girdle. In the leaves the decrease was due mainly to a drop in the level of those compounds that co-chromatographed with the cytokinin glucosides. These compounds were also almost undetectable in the bark above the girdle, where callus was formed. The observed increase in the cytokinin content of the bark above the girdle was due to higher activity in those parts of the chromatograms where zeatin and zeatin riboside occurred. Ringing stimulated the growth of lateral buds below the girdle. These developing buds as well as the bark below the girdle contained very high levels of cytokinins that cochromatographed with zeatin and zeatin riboside.     

The possible involvement of cytokinins in the pathogenicity of Helminthosporium maydis

Green islands/infection sites recorded higher cytokinin activity than surrounding tissue as well as non-inoculated tissue. This activity in infected areas increased with time of incubation while in tissue surrounding the green islands and non-inoculated tissue, cytokinin activity decreased with time of incubation. The culture filtrate extracts of H. maydis had cytokinin activity which increased with growth of the fungus. Cytokinin activity of thin-layer Chromatographic fractions from tissue and culture filtrate extracts revealed that a major portion of the activity was confined to Rf zone 0.6 to 0.8 which co-chromatographed with zeatin and zeatin riboside. Presence of zeatin and zeatin riboside in tissue and culture filtrates was confirmed by high performance liquid chromatography. Cytokinin substances, such as zeatin and zeatin riboside, increase at infection sites with growth of the pathogen suggesting they may be involved in the pathogenicity of H. maydis on maize.     

Phytohormones,Rhizobium mutants,and nodulation in legumes. VII. Identification and quantification of cytokinins in effective and ineffective pea root nodules using radioimmunoassay

Radioimmunoassays (RIA), employing antisera raised in rabbits against bovine serum albumin conjugates of zeatin riboside, dihydrozeatin riboside, and isopentenyladenosine, were used to estimate levels of these cytokinins and their corresponding bases in samples of effective (nitrogen-fixing, Fix⁺), ineffective (nonnitrogen-fixing, Fix^–) pea root nodules and uninoculated roots. Assays were done on extracts of nodule tissue, 1–2 g fresh weight, or approximately 10 g fresh weight of root tissue, and high specific activity [³H]zeatin riboside was added during preparation of the extract for use as a recovery marker. Two different purification procedures were employed, each involving several purification steps. High performance liquid chromatography (HPLC) was the final step in both procedures. Fractions from HPLC were analyzed by RIA using the appropriate antiserum. The cytokinins, zeatin, zeatin riboside, dihydrozeatin riboside, isopentenyl adenine, and isopentenyladenosine were detected and quantified in nodule tissue, and similarly, in root tissue (with the exception of zeatin, which we were unable to quantify in root tissue). Cytokinin levels in nodule tissue were higher than those in root tissue. The major cytokinins detected in nodule tissue were zeatin, followed by zeatin riboside and then dihydrozeatin riboside. The levels of zeatin and zeatin riboside estimated in nodules in the present study by RIA were of the same order of magnitude, though tending to be a little higher, than values obtained previously by bioassay. Dihydrozeatin riboside was identified with confidence for the first time in nodule tissue. There was a general decline with age in cytokinin levels in nodules, but no major qualitative change in nodule cytokinins with age. For theRhizobium strains examined, the data did not indicate a clear correlation between nodule cytokinin levels and the effectiveness of nodules in nitrogen fixation.     

Phytohormones,Rhizobium mutants, and nodulation in legumes. VI. Metabolism of zeatin riboside applied via the tips of nodulated pea roots

[³H]zeatin riboside was supplied in physiological quantities to pea (Pisum sativum L. cv Greenfeast) plants by replacing the root tip with a small vial containing [³H]zeatin riboside, to simulate the normal supply of cytokinin. Radioactivity was transported to the root nodules. Analysis by two-dimensional thin layer chromatography revealed that little³H remained as zeatin riboside in root or nodule tissue at the end of the labeling period (2, 5, or 8 d) and suggested that the following compounds were metabolites of [³H]zeatin riboside: zeatin, adenosine, adenine, the O-glucosides of zeatin and zeatin riboside, nucleotides of adenine and zeatin, and the dihydro-derivatives of many of these compounds.The O-glucosides (and in particular, O--D-glucopyranosyl-9--D-ribofuranosylzeatin) appeared to be more prominent metabolites in the effective nodules formed by strain ANU897 than in the ineffective nodules produced by strain ANU203. However, no other appreciable differences were detected between effective and ineffective nodules in their metabolism of zeatin riboside. There were few marked differences between root and nodule tissue; however, in some experiments, the nodules contained a higher proportion of O-glucoside metabolites, and generally root tissue contained a greater proportion of zeatin and/or dihydro-zeatin, zeatin riboside and/or dihydrozeatin riboside, adenine and the nucleotides of zeatin and adenine, as metabolites.     

The Response of Root Meristems to Colchicine and Indol-3yl-acetic acid in Vicia faba L.

The effects of colchicine and IAA treatments on mitotic activityin various root proliferating tissues have been determined.Lateral root primordia were not affected by IAA, though 24 hfollowing treatment mitotic activity was severely inhibitedin the apical meristems of 1-cm-long attached lateral rootsand primary roots. Primordia were also less sensitive to colchicinetreatment than root apical meristems. Thus telophase figureswere present in the former meristems 3 h following treatment,but not in the latter. Primordia and apical meristems respondedto the same extent, however, to the colchicine-induced increasein number of cells in metaphase, anaphase, and telophase, 3h after treatment began. The apparent difference between largeprimordia and root apical meristems in this respect was dueto the failure of colchicine to penetrate the cells of the formerproliferating tissues as rapidly as the latter. IAA was foundto prevent the increased MI found 24 h following colchicinetreatment only in those meristems where IAA inhibited mitoticactivity at this time. IAA treatments, either alone or withcolchicine, were also found to maintain mitotic activity in1-cm-long lateral roots which were excised from the primaryroots 24 h previously. In such laterals which were not treatedwith IAA, MI was zero at 24 h. It is concluded from the datareported in this paper that, during the development of rootapical meristems, changes take place in the response of cellsto factors affecting mitotic activity.     

Changes in Gene Expression from Diploid to Autotetraploid Status of Lycopersicon esculentum

The cytokinin activity of the root exudate, the leaves, and the apices of vegetative and flowering white lupin plants (Lupinus albus L.) was investigated. The level of cytokinin activity in the root exudate decreased over the 11-week experimental period. Four peaks of cytokinin activity were recorded in the root exudate of 8-week-old plants after fractionation on Sephadex LH-20. Two of these peaks co-eluted with zeatin and zeatin riboside. It is suggested that the remaining peaks represent nucleotide and glucoside cytokinins. The cytokinin levels in extracts of the mature leaves fluctuated slightly over the experimental period. Three peaks of activity co-eluting with zeatin, zeatin riboside and the glucoside cytokinins were recorded in extracts of mature leaves of 8-week-old plants. In the apices cytokinin activity could only be detected in the inflorescences of flowering plants. It would appear that cytokinins produced by the roots accumulate in the fully expanded mature leaves, but are utilized in the rapidly growing apical region and in young expanding leaves.     

Changes in Cytokinin Activity during Seed Germination in Rice (Oryza sativa L.)

Cytokinin activity was determined in dry mature rice seeds,in endosperm and embryo tissues 24, 48 and 72 over imbibitionand in radicles 96 h after germination. Cytokinins with chromatographicproperties similar to zeatin, zeatin riboside, zeatin glucosideand zeatin riboside glucoside were datected in embryo and endosperm,but only the latter two were detected in mature seeds. Cytokininactivity was low during early toges of germination. Qualitativeand quantitative changes in cytokinins were observed in bothembryo and endosperm. The presence of higher cytokinin activityin the endosperm than in the embryo during the first 24 h aftergermination suggests that the endosperm may supply cytokininsuntil the embryo is able to synthaize its own cytokinins. Thepossible significance of high cytokinin glucoside activity inthe embryo early during germination and high cytokinin activityin the radicle during the later stages is discussed. Oryza sativa L., rice, cytokinin, germination, seed     

ANATOMY OF SEEDLING ROOTS OF PSEUDOTSUGA MENZIESII

The seedling root system of Pseudotsuga menziesii (Mirb.) Franco consists of the primary root, active long laterals, long laterals that become mycorrhizal, and short roots that may or may not become mycorrhizal. Numerous adventitious roots arise from the pericycle in young roots and from the vascular cambium and pericycle in older roots following pruning. All actively growing apices have a single plate of initials, a complex zonation of mother cells, and a similar pattern of primary tissue differentiation. Short roots and mycorrhizal short roots have 2 plates of initials, one producing the stele and the other the root cap and cortex, and differentiation occurs close to the apex. Primary and adventitious roots are usually triarch, while long laterals are usually diarch as are all short roots. The latter lack secondary xylem, but mycorrhizal short roots may produce a small amount of secondary phloem.     

Cultivars of Hexaploid Wheat of Contrasting Stature and Chlorophyll Retention Differ in Cytokinin Content and Responsiveness

The work reported here compared cytokinin content and sensitivityin a selection of hexaploid wheat (Triticum aestivum L.) cultivarsusing the following measurements: leaf cytokinins at three timepoints during light-growth and at four 24 h intervals afterlight-grown plants were transferred to darkness; sensitivityof root growth to direct applications of isopentenyl adenosine([9R]iP); and, sensitivity of germination and subsequent rootand shoot growth to 18 h imbibition of seeds in benzyladenine(BA). Accumulation of zeatin riboside-type cultivars was greatestduring light-growth in Tibet Dwarf, a wheat with an extremedwarf phenotype, intermediate in Omar standard and dwarf cultivars,and lowest in the standard and dwarf versions of Itana. Cytokininlevels were otherwise not directly correlated to plant staturein these wheats. There were no cultivar-associated qualitativedifferences in the types of cytokinins detected in this study.During the 16 h light period, the content of zeatin riboside-typecytokinins increased up to tenfold and then declined to basallevels during dark growth. Chlorophyll retention during dark-growthwas correlated with leaf cytokinin content. Data collected ata restricted number of sampling points during dark-growth suggesteda cyclic accumulation of [9R]iP-type cytokinins and the apparentcycle in Tibet Dwarf was offset by 24 h. Tibet Dwarf showedthe greatest root growth inhibition after exposure of seedlingroots to [9R]iP or imbibition of seeds in BA. Neither of thesetreatments affected shoot growth in any of the cultivars. Wheat; Triticum aestivum ; cytokinin; zeatin riboside; benzyladenine; root inhibition     

Changes in the hormonal status of the Taraxacum officinale Web. ovary at early stages of embryogenesis

The hormonal status of the Taraxacum officinale Web. ovary was quantitatively assayed for the first time during early stages of embryogenesis. Apparent concentrations of endogenous cytokinins were measured using two systems of enzyme-linked immunosorbent assay (ELISA). The ELISA systems differed from one another by the specificity for the main endogenous forms of zeatin. The specificity of two heterological ELISA systems based on zeatin- and kinetin-specific antisera was studied. A new immunochemical approach to the problem of differential quantitative determination of natural zeatin forms is suggested. This approach does not require preliminary separation of experimental samples into individual fractions. True concentrations of zeatin and zeatin riboside in the T. officinale ovary were calculated based on the average values of apparent concentrations of endogenous cytokinins. When the embryo sac maturation had been completed, there was a threefold increase in the zeatin riboside concentration within the following 12 h. By the time of the first division of an unfertilized ovicell (i.e., within the next 12 h), there had been a twofold decrease in the zeatin riboside concentration. Therefore, at early stages of division of the unfertilized ovicell the zeatin riboside concentration virtually returned to the initial level. In contrast to zeatin riboside, there was a steady trend toward an increase in the zeatin concentration in the T. officinale ovary. Within the first 12 h and the next 12 h after completion of the embryo sac maturation, the zeatin concentration was increased 1.5-fold and 2-fold, respectively. The results of this work provide a pioneering insight into the dynamics of various natural forms of zeatin during the reproductive process. The immunochemical approach to quantitative monitoring of various natural forms of zeatin and their dynamics during embryogenesis suggested in this work can be extended to similar biological, medical, and agricultural problems of differential determination of low-molecular-weight agents of similar structure but different biological activity.     

Root Development and Source-Sink Relations in Carrot, Daucus carota L: II. EFFECTS OF ROOT PRUNING ON CARBON ASSIMILATION AND THE PARTITIONING OF ASSIMILATES

Physiological responses to root pruning were investigated bycomparing ¹⁴CO₂ fixation rates, the partitioning of ¹⁴C-labelledassimilate, and soluble and insoluble carbohydrate levels inthe leaves of carrot plants following the removal of some ofthe fibrous roots, or fibrous roots and part of the tap root.Root pruning reduced ¹⁴CO₂ fixation by 28–45% but leafspecific activity (¹⁴C assimilation g-¹ leaf fresh weight) wasunchanged. The proportion of total assimilate exported to theroot system increased following root pruning and this was atthe expense of the developing leaves. In younger plants (wherethe tap root received 10% of the assimilate) the supply of ¹⁴Cto the tap root was maintained in spite of root pruning. However,shortening the tap root to 3 cm in older plants (in which 30%of the fixed 14C was normally exported to the developing storageorgan), reduced its sink capacity and resulted in slightly greaterretention of ¹⁴C in the mature leaves. Greater concentrationsof insoluble carbohydrate were found in the mature leaves followingroot pruning but soluble sugar content was unaffected. Onlysmall differences were observed in the distribution of ¹⁴C betweensoluble and insoluble carbohydrate fractions when plants werefed ¹⁴CO₂ several days after the root pruning operations. Thesephysiological responses were mainly associated with the removalof fibrous roots and support the view that the fibrous rootsystem is more important than the developing storage organ inregulating growth in young carrot plants.     

Rapid increases in cytokinin concentration in lateral buds of chickpea (Cicer arietinum L.) during release of apical dominance

We examined the role of cytokinins (CKs) in release of apical dominance in lateral buds of chickpea (Cicer arietinum L.). Shoot decapitation or application of CKs (benzyladenine, zeatin or dihydrozeatin) stimulated rapid bud growth. Time-lapse video recording revealed growth initiation within 2 h of application of 200 pmol benzyladenine or within 3 h of decapitation. Endogenous CK content in buds changed little in the first 2 h after shoot decapitation, but significantly increased by 6 h, somewhat later than the initiation of bud growth. The main elevated CK was zeatin riboside, whose content per bud increased 7-fold by 6 h and 25-fold by 24 h. Lesser changes were found in amounts of zeatin and isopentenyl adenine CKs. We have yet to distinguish whether these CKs are imported from the roots via the xylem stream or are synthesised in situ in the buds, but CKs may be part of an endogenous signal involved in lateral bud growth stimulation following shoot decapitation. To our knowledge, this is the first detailed report of CK levels in buds themselves during release of apical dominance. Received: 12 December 1996 / Accepted: 7 January 1997     

Plant-water relations in wheat as influenced by root pruning

Summary A greenhouse study in which 24, 54 and 71 per cent roots of wheat (Triticum aestivum L.) were pruned on the 73rd day from the date of planting (anthesis stage) showed that during a 7-day period following root pruning, total transpiration and leaf water potential were significantly lower (P=0.05) and the stomatal resistance was significantly higher (P=0.05) where 54 and 71 per cent roots were pruned, as compared to no root pruning or 24 per cent root pruning. The leaf relative water content, however, showed no significant differences. Thus about one-fourth root sytem could be reduced without adversely affecting the plant-water status.     

Changes in Phytohormone Status in Stems and Roots after Decapitation of Pea Seedlings

Experiments were performed on the first and second internodes and 4-cm-long apical segments of main roots of pea (Pisum sativum L.) seedlings, grown in the light and decapitated above the second node on the seventh day after seed germination. Endogenous phytohormones were measured by the enzyme-linked immunosorbent assay during three days after decapitation of seedlings. The IAA level in the internodes decreased 2–3 times on the second day after decapitation of seedlings while the cytokinin level increased 5–6 times for zeatin and zeatin riboside (Z and ZR) and 1.5–2 times for isopentenyl adenine and isopentenyl adenosine (IP and IPA). In contrast to internodes, the IP and IPA contents in the roots of decapitated seedlings did not change, but the levels of Z and ZR increased 1.5–2 times compared to intact plant roots. The IAA level in the apical region of root remained almost unchanged after the removal of shoot apex. It was concluded that the apical meristem of the main root is not the site of the cytokinin response to the auxin signal coming from the stem apex and that a slight accumulation of Z and ZR after decapitation is due to upper zones of the root. There was no difference in the content of gibberellin-like substances between the internodes of intact and decapitated seedlings. However, the content of gibberellins (GA) in the root tip decreased after decapitation of seedling, which suggests an essential role of apical bud in supplying the root with GA and/or intermediates for their biosynthesis.