Inverse Correlation between ABA Content and Germinability throughout the Maturation and the In Vitro Culture of the Embryo of Phaseolus vulgaris

Prevost, I. and Le Page-Degivry, M. Th. 1985. Inverse correlationbetween ABA content and germinability throughout the maturationand the in vitro culture of the embryo of Phaseolus vulgaris.—J.exp. Bot. 36: 1457–1464. Changes in embryo abscisic acid (ABA) content during the maturationof the seed of Phaseolus vulgaris cv. Contender were followed,using a radio-immunoassay. The pattern of change is similarto that already described in several species: a rapid increase(from the 18th to 29th day after anthesis), was followed bya decrease, the ABA level being ten times lower on the 48thday than on the 29th. Embryos isolated from the 18th to the48th day after anthesis were able to ‘germinate’when cultivated on a mineral medium supplemented with sucroseand agar. The development pattern varied throughout the embryogenesisand could be correlated with the differentiation of the embryoat the time of isolation. Before germination could take place,we observed a lag phase, the duration of which could be correlatedwith embryo ABA content. As ABA content increased in the youngestembryos the duration of the lag phase increased. In the sameway, the number of days to germination was shown to diminishas ABA content decreased. Inverse correlation between ABA contentand germinability was thus demonstrated throughout the developmentof the embryo. During in vitro culture, free ABA content decreased in the embryoand reached low values a few days before germination occurred.So the beginning of root elongation in culture was again wellcorrelated with the disappearance of free endogenous ABA. Atransfer experiment inducing an earlier germination associatedwith a more limited development suggests that the lag phaseis associated with an active continuation of embryonic development Key words: Embryo maturation, abscisic acid, germinability     

Maturation and germination of Phaseolus vulgaris embryonic axes in culture

In this study of embryo development in Phaseolus vulgaris L., we found that immature embryonic axes placed in culture show a growth lag before germinating. The length of this lag phase varies according to axis age at excision, but is not affected by transfer to fresh medium, alteration of sucrose concentration between 0.5 and 2%, or whether the culture medium is liquid or agar-solidified. The lag phase was shortened by both actinomycin D and cordycepin treatment, and by treatment with 10^-5 to 10^-6 M benzyladenine. The effect of abscisic acid (ABA) varied with concentration: below a certain level, it had no effect on the lag phase, but above that level it inhibited, germination. This threshold concentration was 10^-7 M for 20-d-old axes but increased to 10^-5 M by the time the axes were 32 to 34 d old. To determine whether the axes were continuing their embryonic development during the lag phase, we tested them for desiccation-tolerance and for synthesis of phaseolin, a seed storage protein which is specific for embryos of P. vulgaris. The ability to germinate after rapid desiccation was acquired by axes at 26 d past anthesis; when axes younger than this were placed in culture, they developed desiccation-tolerance during the lag phase of growth, indicating that they were continuing embryonic maturation. Phaseolin was present in isolated axes, although at lower levels than in cotyledons. It accumulated during axis development in parallel with total protein, staying at about 1% of total protein content. When isolated immature axes were pulsed with ³H-or ¹⁴C-amino acids, they incorporated label into phaseolin, shown by precipitation with anti-phaseolin antibody. Isolated axes from mature seeds, however, did not synthesize detectable amounts of phaseolin. Immature axes cultured in vitro for a period of one to several days continued synthesizing phaseolin until the day prior to visible germination. Treatment of cultured axes with ABA increased the amount of precursor amino acids incorporated into protein, but had a small or no effect on the relative proportion of phaseolin synthesized. We conclude that P. vulgaris axes in culture continue to develop embryonically for a period of time which seems to be under intrinisc control by the axis. This contrasts with precocious germanation, a pattern of embryo behavior seen in many other species. When such embryos are excised from seeds while immature and placed in culture, they switch promptly from embryo development into germination. If ABA or water stress is responsible for preventing precocious germination, it may be that a high level of ABA is maintained or synthesized internally by embryonic axes of Phaseolus, while in other embryos the maternal environment supplies ABA and/or causes water stress.Abbreviations ABA Abscisic acid - BA benzyladenine     

A promotive action of kinetin on amylase activity in cotyledons of Phaseolus vulgaris

Kinetin and the embryo axis acted similarly in bringing abouta promotion of amylase activity in cotyledons of Phaseolus vulgaris.No promotive effect of gibberellic acid or indole-3-acetic acidon amylase activity could be detected. It is suggested thatthe regulatory action of the embryo axis on starch degradationin the cotyledons of P. vulgaris is mediated by cytokinins. (Received May 4, 1970; )     

Response of Cultured Embryos of Phaseolus vulgaris and Hordeum vulgare to Farnesol

Excised embryos of Phaseolus vulgaris incubated in a mediumcontaining 10 mg dm^–3 farnesol showed enhanced root growthwhereas the leaves remained rudimentary At lower concentrationsof exogenous farnesol normal leaf development occurred and rootgrowth was comparable to untreated cultures. Enhanced root growthalso occurred when excised embryos of Hordeum vulgare were treatedwith farnesol but only at 10 mg dm^–3 and this treatmentdid not prevent leaf growth X-ray micro-probe analysis of leavesrevealed an increased phosphorus content in P vulgaris and adecreased sulphur content in H vulgare in comparison to untreatedplants. Hordeum vulgare L., barley, Phaseolus vulgaris, bean, embryo culture, farnesol, X-ray microprobe analysis, root growth     

The Role of ABA in the Control of Apple Seed Dormancy Re-appraised by Combined Gas Chromatography-Mass Spectrometry

Measured by GC—MS²—SIR³, endogenous ABA⁴ in embryonicaxes of seeds of Malus pumila L. cv. Golden Delicious decreased8-fold and cotyledon ABA by only 60%, during 10–50 d ofstratification at 5 ?C, after ABA leaching during an initial24 h soaking. During stratification, the percentage germinationof embryos transferred to 17?C showed a significant linear dependenceon log_e of ABA levels in the axes at transfer. Between 50 and70 d, ABA levels increased markedly in axes and testa both ofstratified seeds and seeds allowed to re-dry at 17 ?C afterinitial soaking. The ability of fully stratified axes with elevatedendogenous ABA to germinate indicated that stratification haddecreased their ABA sensitivity. Changes in cotyledon ABA couldnot account for the promotory effect of cotyledons on germinationduring the first 30 d of stratification. Loss of testa inhibitionof germination during stratification was not linked with changesin testa ABA. Stratification markedly increased the sequestrationin the axes of exogenous ABA supplied via the cotyledons. Changesboth in axis ABA levels and sensitivity were thus correlatedwith dormancy release, but subject to modifying control by thecotyledons and testa not involving ABA. Rehydration of driedseeds affected axis ABA later during storage via mechanismsunconnected with dormancy. Key words: ABA, seed dormancy, stratification     

The Utilization of Cotyledonary Reserves in Phaseolus vulgaris L. cv. Carioca: II. THE EFFECTS OF 6-BENZY LADENINE AND GIBBERELLIC ACID UPON EMBRYONATED AND DETACHED COTYLEDONS

The presence of the embryo axis promotes starch hydrolysis incotyledons of Phaseolus vulgaris and, although reduced ratesof enzyme activity proceed in its absence, weight loss fromthe cotyledon is only significant in its presence. Applicationof gibberellic acid at various concentrations had no effectupon amylase or protease activity in either detached or embryonatedcotyledons: this was not the case for 6-benzyladenine whichpromoted both. In detached cotyledons amylolytic activity wasdirectly proportional to the concentration of applied 6-BA;concentrations below 10^–6 M were unable to completelysubstitute for the presence of the axis, while, above 10^–6M, hydrolytic activity higher than that of treated embryonatedcotyledons was observed. Increasing the concentration of 6-BAhad no effect, however, upon the amylolytic activity of embryonatedcotyledons. Detached cotyledons showed an increased chlorophyllcontent and 6-BA treatment further increased this; the proportionsof chlorophyll a and chlorophyll b remained unaltered. An inhibitor of amylase activity, destroyed by heating, is presentin embryonated cotyledons and can depress amylolysis in detachedcotyledons which appear to be inhibitor-free. The results intotal suggest that the developing axis may regulate reservehydrolysis in three distinct manners: firstly, by stimulatingthe synthesis of amylase via hormonal control, most probablya cytokinin; secondly, by checking the rate of amylolysis bymeans of an amylase inhibitor; and, thirdly, by being a sinkfor the products of reserve degradation.     

Seed Dormancy in Acer: The Role of Testa-imposed and Embryo Dormancy in Acer velutinum

The embryo dormancy shown in freshly harvested samples of Acervelutinum seeds is weakly established and very short-lived.Loss of this embryo dormancy occurred during post-harvest fruitstorage at either 5 or 17 C. In contrast, the dormancy of intactfruits and seeds was overcome only during storage at the lowertemperature. Removal of the cotyledons from embryos of freshlyharvested fruits allowed more rapid germination of the embryonicaxes, indicating that the cotyledons exert an inhibitory effect,although the axes still retained a measure of innate dormancy.The inhibitory effect of the cotyledons became less marked withincreasing duration of fruit storage, this loss of inhibitoryeffect occurring at both storage temperatures. Applied ABA stronglysuppressed germinative capacity in intact embryos and isolatedembryonic axes from freshly harvested fruits, but when ABA wasapplied to embryos of fruits that had been stored for variousperiods at 5 or 17 C, the inhibitory effect was first weakenedand then lost with increased storage. Although dormancy in the seeds of A. velutinum may be describedas intermediate between testa-imposed dormancy and true dormancy,it is perhaps more properly included in the former category. Acer velutinum Boiss. var. vanvolxemii, abscisic acid, embryo dormancy, germination, seed storage, testa-imposed dormancy, tissue sensitivity     

cytokinin-induced amylolytic activity in bean cotyledons: Identification of the regulated enzyme

Separation of an extract of cotyledons of Phaseolus vulgarison a column of Sephadex G-200 revealed three amylolytic fractions.The slower migrating fraction hydrolyzed/ß-Limit dextrinazure and was inhibited by EDTA. The activity of this fractionwas enhanced by the embryo axis and this effect could be fullyreplaced by kinetin or benzyladenine. These results suggestthat the bean embryo axis exerts a promotive influence on theactivity of a-amylase in the cotyledons and that this effectis mediated by cytokinins. The other two amylolytic fractions did not show a-amylase activity.No effect of the embryo axis or of cytokinins on their activitycould be noted. ¹Present address: The Thimann Laboratories, University of California,Santa Cruz, California, U.S.A. (Received June 27, 1979; )     

Cytokinins in Germinating Seeds of Phaseolus vulgaris L. II. Transport and Metabolism of 8[14C]t-Zeatin Applied to the Radicle

The application of 8[¹⁴C]t-zeatin to the radicle tips of germinatingPhaseolus vulgaris seeds revealed that cytokinins are transportedrapidly from the embryonic axis to the cotyledons, and are utilizedand metabolized extensively in these organs. The informationobtained on the transportation between the different parts ofthe embryo is consistent with the view that the mobilizationof food reserves from the cotyledons is controlled by cytokininswhich originate in the embryonic axis. Tentative identificationof the radioactive metabolites formed indicate that the appliedzeatin was altered by side-chain cleavage and by substitutionto the adenine ring. Phaseolus vulgaris, bean, germination, cytokinins, transport, radicle     

Inhibition of Light-Stimulated Leaf Expansion by Abscisic Acid

Abscisic acid (ABA) applied to intact bean (Phaseolus vulgaris)leaves or to isolated leaf discs inhibits light-stimulated cellenlargement This effect may be obtained with 10^–4 molm^–3 ABA, but is more significant at higher concentrations.The inhibition of disc expansion by ABA is greater for discsprovided with an external supply of sucrose than for discs providedwith KC1, and may be completely overcome by increasing the KC1concentration externally to 50 mol m^–3. Decreased growthrate of ABA-treated tissue is not correlated with loss of solutesfrom growing cells, but is correlated with a decrease in cellwall extensibility. ABA does not prevent light-stimulated acidificationof the leaf surface, and stimulates the acidification of theexternal solution by leaf pieces. However, the capacity of thecell walls to undergo acid-induced wall loosening is diminishedby ABA-treatment. The possibility that ABA acts directly byinhibiting growth processes at the cellular level, or indirectlyby causing stomatal closure, is discussed. Key words: Phaseolus vulgaris, ABA, Inhibition, Leaf expansion     

Normalizing Development of Cultured Triticum aestivum L. Embryos: I. LOW OXYGEN TENSIONS AND EXOGENOUS ABA

Wheat (Triticum aestivum L.) embryos form in dynamically-regulatedovular environments. Our objectives were to improve developmentof cultured immature wheat embryos by simulating, in vitro,abscisic acid (ABA) levels and O₂ tensions as found in wheatovules during zygotic embryogenesis. We characterized from intactwheat kernels embryo respiration, embryo morphology and embryoand endosperm + ABA levels at 13, 19 and 25 d post-anthesis(DPA). Young (13 DPA) embryos were then excised and culturedin vitro, where they were exposed to 0·2 or 2·Ommol m^–3 ±ABA and 2.·1, 2·5 or 7·4mol m^–3 (6, 7 and 21%, respectively) gaseous O₂. At 6and 12 d in culture, + ABA levels, embryo respiration and embryomorphology were characterized by treatment. Thirteen-day-oldembryos from two different plant populations differed by 17-foldin initial ABA content. However, this difference did not affectprecocious germination in vitro, nor did it affect the amountof exogenous ABA required to reduce precocious germination by40%. In this respect, embryos from both populations were equallysensitive to exogenous ABA. Cavity sap O₂ levels (2·1to 2·5 mol m^–3) were much more effective in preventingprecocious germination of cultured embryos than were cavitysap levels of ABA (0·2 to 2·0 mmol m^–3).The combination of physiological levels of both ABA and O₂ largelynormalized DW accumulation and embryo morphology without alteringendogenous + ABA levels. Residual respiration of cultured embryoswas higher than that of embryos grown in situ, and was not influencedby the exogenous O₂ and ABA treatments Key words: Abscisic acid, embryo development, oxygen tensions, respiration, wheat     

Correlative Inhibition of Lateral Bud Growth in Pisum sativum L. and Phaseolus vulgaris L.: Studies of the Role of Abscisic Acid

The possibility has been investigated that abscisic acid (ABA)might act as a correlative inhibitor of lateral bud growth inPisum sativum and Phaseolus vulgaris. Application of ABA insmall quantities (2µg) to axillary buds on decapitatedplants of P. sativum caused appreciable inhibition of theirgrowth, and induced a compensatory growth of the bud on an adjacentnode. Application of this same quantity of ABA to axillary budson decapitated plants of Phaseolus vulgaris was without effect,but a high concentration in lanolin (1 mg g^–1) did substantiallyreduce bud outgrowth. Endogenous ABA-like substances in Phaseolusvulgaris, detected by bioassay and electron capture g.l.c.,were present in similar concentrations in shoot tips, lateralbuds on intact plants and lateral buds on plants decapitated24 h earlier. The effects of applied ABA suggested that it might be involvedin the mechanism of correlative inhibition in Pisum sativum,but it was not possible to test this hypothesis by determiningendogenous ABA levels in axillary buds because of their smallsize. The evidence presented here suggests that ABA is not acorrelative inhibitor in Phaseolus vulgaris even though at highconcentration it can inhibit the growth of axillary buds.     

Gibberellins in the embryonic axes of tall and dwarf beans and their changes with initial growth

Ten gibberellin-like activities were detected in the dry embryonicaxes of tall (cv. Kentucky Wonder) and dwarf (cv. Masterpiece)beans (Phaseolus vulgaris L.) using the lettuce hypocotyl assayof thin-layer chromatograms; 2 in the non-acidic ethyl acetatefraction (NEI, NEII), 3 in the acidic ethyl acetate fraction(AEI, AEII, AEIII), 2 in the non-acidic n-butanol fraction (NBI,NBII) and 3 in the acidic n-butanol fraction (ABI, ABII, ABIII).There was no qualitative difference in these gibberellins betweenthe tall and dwarf axes, but all, particularly AEIII, NBII andABIII as the main gibberellins in the axes, were contained muchmore abundantly in the tall axes. In both axes the gibberellinactivities of most fractions decreased during germination.Theamounts of some gibberellins in tall axes without cotyledonswere greater than those in axes with cotyledons at 48–72hr of germination. Neither AMO-1618 nor CCC caused significantreduction in the levels of the gibberellins. Axis growth inthe early germinating period depended on the gibberellins storedin the axis, itself. (Received November 26, 1974; )     

Cytokinins in Germinating Seeds of Phaseolus vulgaris L. III. Transport and Metabolism of 8[14C]t-Zeatin Applied to the Radicle

The application of 8[¹⁴C]t-zeatin to the cotyledons of germinatingbean seeds demonstrated that cytokinins are not readily exportedfrom the cotyledons to the embryonic axis during the early stagesof this process. In the cotyledons the applied zeatin is metabolizedextensively to metabolites which are polar and which occur atR_F 0·2–0·5 on paper chromatograms. Thesemetabolites are stable and are not readily exported from thecotyledons. In contrast the metabolites found at R_F 0–0·2are more readily exported. When exported to the radicles andplumules a large proportion of the translocated metaboliteswere converted to compounds which on paper co-chromatographedwith zeatin. This seems to suggest that the embryonic axis hasthe capacity to synthesize cytokinins and that some of the metabolitesformed during its catabolism can also be used for its synthesis. Phaseolus vulgaris, bean, germination, cytokinins, transport, cotyledons     

Absorption, Distribution, Metabolism and Leaching of [14C] ABA during Culture of Apple Embryos

Barthe, Ph. and Bulard, C. 1987. Absorption, distribution, metabolismand leaching of [¹⁴C] ABA during culture of apple embryos.—J.exp. Bot. 38: 1002–1011. It has been known for some time that dormant embryos, laid flaton damp filter-paper one cotyledon only being in direct contactwith it (C/2M mode of culture), exhibit unequal growth and greeningof their cotyledons. The aim of this work was to investigatewhether this particular mode of culture led to detectable differencesbetween the two cotyledons in the distribution and metabolismof [¹⁴C] abscisic acid (ABA). Two different approaches wereused, the material in both cases being dormant embryos of Pyrusmalus L. cv. Golden Delicious cultured at 23°C in darkness.In a first experiment, the embryos were cultured directly inthe C/2M mode and in the presence of 10^–2 mol m^–3[¹⁴C] ABA. In these conditions marked differences in the distributionof radioactivity between the lower (LC) and upper (UC) cotyledonappear after 24 h of culture. After 5 d, amounts of total radioactivitywere four times higher in LC than in UC, and the level of ABAwas three times higher. Metabolism was extremely active in LC,since certain metabolites were found in relative percentages(percentages with respect to the total radioactivity of thecotyledon) equivalent to (esters, glucosides), or even higher(dihydrophaseic acid, DPA) than those found in UC. It is suggestedthat this high level of metabolism in LC limits the availabilityof the transportable molecule, that is to say ABA. In a secondexperiment double culturing was carried out. The embryos werefirst cultured in the presence of 10^–2 mol m^–3 [¹⁴C]ABA in conditions which ensured equal distribution of ABA andits metabolites between the two cotyledons. After 5 d they weretransferred to an ABA-free medium and cultured in the C/2M mode.After 2 d and 5 d of culture, dissymmetry between the two cotyledonswas again noted; UC this time containing greater amounts ofradioactivity than LC. The differences, however, were less markedthan in the first experiment. This dissymmetry is due to leachingof a certain amount of radioactivity from LC, which is onlypartially compensated for by migration from UC. Key words: ¹⁴C-ABA distribution, leaching and metabolism, embryo culture, embryo dormancy     

Cytokinins in Germinating Seeds of Phaseolus vulgaris L. I. Changes in Endogenous Levels Within the Cotyledons

Ethanolic extracts from the cotyledons of mature dry Phaseolusvulgaris L. seed yielded cytokinin-like activity which co-chromatographedwith zeatin and ribosylzeatin. Under conditions which stimulatedgermination and cotyledon expansion, the level of these cytokininsdecreased rapidly in both intact embryos and excised cotyledons.In the excised cotyledons the decrease was continuous, resultingin very low levels of cytokinin being detected after 4 daysof incubation. With the embryonic axis present, however, theinitial decrease was arrested and reversed after 3 days. Thissuggests that the cotyledons do not synthesize cytokinins butthat these hormones are imported from the embryonic axis, particularlyonce radicle growth is well under way. Phaseolus vulgaris, bean, cotyledons, cytokinins, germination     

The Role of Maturation Drying in the Transition from Seed Development to Germination: VII. EFFECTS OF PARTIAL AND COMPLETE DESICCATION ON ABSCISIC ACID LEVELS AND SENSITIVITY IN RICINUS COMMUNIS L. SEEDS

During mid-development (25–40 d after pollination: DAP)of the castor bean seed the amount of abscisic acid (ABA) increasesin both the endosperm and the embryo, declining substantiallythereafter until there is little present in the mature dry (60DAP) seed. Premature desiccation of the seed at 35 DAP alsoleads to a major decline in ABA within the embryo and endosperm.Partial water loss from the seed at 35 DAP which, like naturaland premature desiccation, leads to subsequent germination uponreturn of the seed to full hydration, causes a much smallerdecline in ABA levels. In contrast, ABA declines substantiallyin the non-dried (hydrated) control at 35 DAP, but the seedsdo not germinate. Hence, a clear negative correlation betweenABA content and germinability is not observed. Both drying,whether natural or imposed prematurely, and partial drying decreasethe sensitivity of the isolated embryo to exogenous ABA by about10-fold. The protein synthetic response of the castor bean embryo exposedto 0.1 mol m^–3 ABA following premature desiccation exhibitssome similarity to the response of the non-dried developingembryo—in both cases the synthesis of some developmentalproteins is enhanced by ABA, and germination is suppressed.Germination of mature seeds is also suppressed by 0.1 mol m^–3ABA, but the same developmental proteins are not synthesized.In the cotyledons of prematurely-desiccated seed, some proteinsare hydrolysed upon imbibition in 0.1 mol m^–3 ABA, a phenomenonthat occurs also in the cotyledons of similarly treated matureembryos, but not in developing non-dried embryos. Hence theembryo exhibits an ‘intermediate’ response uponrehydration in 0.1 mol m^–3 ABA following premature desiccation;viz. some of the responses are developmental and some germinative.Following natural or imposed drying, the isolated embryo becomesrelatively insensitive to 0.01 mol m^–3 ABA: germinationis elicited and post-germinative reserve breakdown occurs inthe radicle and cotyledons. The reduced sensitivity of the embryoto ABA as a consequence of desiccation may be an important factorin eliciting the switch to germination and growth within thewhole seed. Key words: Abscisic acid, desiccation, astor bean endosperm, seed development, germination, protein synthesis, isolated embryos, hormone sensitivity     

Apical Dominance in Phaseolus vulgaris L.

Although determinations of the ABA content of lateral buds ofPhaseolus vulgaris revealed no difference between decapitatedand intact control plants in the first 12 h following decapitation,a relative decrease in the ABA content of lateral buds of decapitatedplants was detectable 24 h following decapitation. Shoot decapitationwas also observed to result in a decrease in the ABA contentof stem tissue. The application of IAA to the stem of decapitatedplants prevented these changes and increased the ABA contentof stem tissue relative to that of intact plants. The levelsof IAA and ABA were also determined in the stem tissue fromthe nodes of intact bean plants. The possible interdependenceof these two plant hormones was further investigated by a studyof [2_–14ClABA metabolism. The results are discussed inrelation to the possible role of these hormones in apical dominance. Key words: Apical dominance, Abscisic acid, Indole-3-acetic acid     

Uptake of [2-14C]abscisic acid and distribution of 14C in apple embryos

Pyrus malus L. var. Golden delicious embryos were incubated with (±)-[2-¹⁴C]abscisic acid (ABA) (10^-5 M, 355 kBq mol^-1). After incubations of various durations, the radioactivity was measured in whole embryos, cotyledons, and embryonic axes.With either 48-h or 16-d incubation periods, the uptake of [¹⁴C]ABA depended upon the mode of culture used. The lowest values corresponded to the absorption by the embryonic axis, the highest to the absorption by the distal parts of the two cotyledons. The cotyledons accumulated the main part of the radioactivity under all conditions. Dormant and almost completely after-ripened embryos cultivated for 4 d showed no significant differences in the radioactivity uptake for identical modes of culture. There was a linear relationship between exogenous ABA concentrations (0.5 to 3.10^-5 M) and ABA uptake for embryos cultivated for 4 d with the distal parts of the cotyledons immersed in the medium.Abbreviations ABA abscisic acid. RM, RM⁺, C/2 M, and CM are different modes of embryo cultures: embryonic axis immersed alone (RM), together with the proximal parts of the cotyledons (RM⁺); distal parts of the cotyledons immersed alone (CM); embroyo flat on the medium, the root and the external surface of one cotyledon being in contact with the medium (C/2 M) - PP proximal parts of the cotyledons - DP distal parts of the cotyledons     

Internodal Extension in the First Trifoliate Leaf Axillary Bud of Phaseolus vulgaris L. following Shoot Decapitation

Phaseolus vulgaris L. decapitated at the third internode showedaccelerated growth of the uppermost axillary bud remaining onthe stem (the first trifoliate axillary bud) after a lag periodof 3–5 h Much of the initial growth increment could beattributed to cell expansion Phaseolus vulgaris L, dwarf bean, correlative inhibition, cell expansion